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Full text of "Annals of the South African Museum. Annale van die Suid-Afrikaanse Museum"

w' 



HARVARD UNIVERSITY 

Library of the 

Museum of 

Comparative Zoology 



ANNALS OF THE ANNALE VAN DIE 

SOUTH AFRICAN MUSEUM SUID-AFRIKAANSE MUSEUM 



VOLUME 74 BAND 74 



ANNALS OF THE SOUTH AFRICAN MUSEUM 
ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM 

VOLUME 74 BAND 




THE TRUSTEES OF THE 
SOUTH AFRICAN MUSEUM 
CAPE TOWN 



DIE TRUSTEES VAN DIE 

SUID-AFRIKAANSE MUSEUM 

KAAPSTAD 



1977-1978 



524 ** 



§ SET, PRINTED AND BOUND IN THE REPUBLIC OF SOUTH AFRICA BY 
THE RUSTICA PRESS (PTY.) LTD., WYNBERG, CAPE 



LIST OF CONTENTS 



Page 
Avery, D. M. 

Past and present distribution of some rodent and insectivore species in the southern 

Cape Province, South Africa: new information. (Published December 1977.) 201 

Eastwood, E. B. 

Notes on the scorpion fauna of the Cape. Part 3. Some observations on the distri- 
bution and biology of scorpions on Table Mountain. (Published January 1978.) 229 

Eastwood, E. B. 

Notes on the scorpion fauna of the Cape. Part 4. The burrowing activities of some 

scorpionids and buthids (Arachnida, Scorpionida). (Published January 1978.) 249 

Glaessner, M. F. 

Re-examination of Archaeichnium, a fossil from the Nama Group. (Published April 

1978.) 335 

Griffiths, C. 

The South African Museum's Meiring Naude cruises. Part 6. Amphipoda. (Published 

November 1977.) . . 105 

Grindley, J. R. 

A new species of Tortanus (Crustacea, Copepoda) from South Africa. (Published 

January 1978.) 219 

Kennedy, W. J. & Klinger, H. C 

Cretaceous faunas from Zululand and Natal, South Africa. A Jauberticeras from 

the Mzinene Formation (Albian). (Published November 1977.) . . . . 1 

Kennedy, W. J. & Klinger, H. C. 

Cretaceous faunas from Zululand and Natal, South Africa. A Flickia from the 

Cenomanian of northern Zululand. (Published January 1978.) . . . . 211 

Kennedy, W. J. & Klinger, H. C. 

Cretaceous faunas from Zululand and Natal, South Africa. The ammonite family 

Lytoceratidae Neumayr, 1875. (Published February 1978.) 257 

Kensley, B. 

The South African Museum's Meiring Naude cruises. Part 5. Crustacea, Decapoda, 

Reptantia and Natantia. (Published December 1977.) 13 

Kensley, B. 

The South African Museum's Meiring Naude cruises. Part 7. Marine Isopoda. 

(Published January 1978.) 125 

Klinger, H. C. see Kennedy, W. J. 

McKenzie, K. G. 

Illustrated generic key to South African continental Ostracoda. (Published Novem- 
ber 1977.) . . . . 45 

Millard, N. A. H. 

The geographical distribution of southern African hydroids. (Published January 

1978.) 159 



NEW GENERIC NAMES PROPOSED IN THIS VOLUME 





Page 


lzinkala Griffiths, 1977 


115 


Kapcypridopsis McKenzie, 1977 


46 


Sarscypridopsis McKenzie, 1 977 


48 


Spinarcturus Kensley, 1978 


135 


Valet tiella Griffiths, 1977 


116 



SOU 7SU^ 

VOLUME 74 PART 1 NOVEMBER 1977 



ISSN 0303-2515 



MU9. COMP. ZOOL. 

LIBRARY 

dAN 3 1 1978 

HARVARD 
UNIVERSITY 



ANNALS 




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Bullough, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan. 

Fischer, P.-H. 1948. Donnees sur la resistance et de le vitalite des mollusques. /. Conch., Paris 88: 100-140. 

Fischer, P.-H., Duval, M. & Raffy, A. 1933. Etudes sur les echanges respiratoires des littorines. Archs 

Zool. exp. gen. 74: 627-634. 
Kohn, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the TrincOmalee region of Ceylon. 

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Kohn, A. J. 19606. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean. 

Bull. Bingham oceanogr. Coll. 17 (4): 1-51. 
Thiele, J. 1910. Mollusca: B. Pblyplacophora, Gastropoda marina, Bivalvia. In: Schultze, L. Zoologische 

und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Sud-Afrika 4: 269-270. 

Jena: Fischer. Denkschr. med.-naturw. Ges. Jena 16: 269-270. 

{continued inside back cover) 



ANNALS OF THE SOUTH AFRICAN MUSEUM 
ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM 



Volume 74 Band 
November 1977 November 
Part 1 Deel 




CRETACEOUS FAUNAS FROM ZULULAND 

AND NATAL, SOUTH AFRICA 

A JAUBERTICERAS FROM THE MZINENE 

FORMATION (ALBIAN) 

By 

WILLIAM JAMES KENNEDY 

& 
HERBERT CHRISTIAN KLINGER 



Cape Town Kaapstad 



The ANNALS OF THE SOUTH AFRICAN MUSEUM 

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CRETACEOUS FAUNAS FROM ZULULAND AND NATAL, 

SOUTH AFRICA 
A JAUBERTICERAS FROM THE MZINENE FORMATION (ALBIAN) 

By 

William James Kennedy 

Geological Collections, University Museum, Oxford 

& 

Herbert Christian Klinger 

South African Museum, Cape Town 

(With 6 figures) 
[MS. accepted 28 April 1977] 

ABSTRACT 

The ammonite subfamily Gabbioceratinae Breistroffer, 1953, is represented in the 
Cretaceous of southern Africa by a unique specimen of Jauberticeras collignoni Murphy from 
the Albian Mzinene Formation exposed along the Mzinene River, Zululand. The specimen 
shows details of suture, lirae and constrictions not previously known in the species, until now 
based on the unique Madagascan holotype alone, allowing a fuller diagnosis, description and 
comparison with other species. The holotype and the types of the closely related species 
J. jaubertianum (d'Orbigny) and J. vilhittreysi Murphy are reillustrated for comparative 
purposes. 



CONTENTS 




Introduction .... 

Systematic palaeontology 

Acknowledgements 

References .... 


PAGE 

1 
2 

11 

11 



INTRODUCTION 

In two previous papers (Kennedy & Klinger 1977, and in press) the authors 
have documented the representatives of the great majority of the ammonite 
superfamily Tetragonitaceae Hyatt, 1900 known from the Cretaceous sediments 
of Zululand and Natal. In the course of examining the collections of the late 
Dr E. C. N. van Hoepen, recently transferred to the South African Museum 
from the National Museum Bloemfontein, one of the authors (H. C. K.) recog- 
nized a further representative of this superfamily, in the form of a Jauberticeras 
Jacob, 1907, from the Upper Albian Mzinene Formation of Zululand. 

Jauberticeras is one of the two genera referred to the subfamily Gabbio- 
ceratinae Breistroffer, 1953 (the other is the genus Gabbioceras Hyatt, 1900); 
the specimen represents not only the first record of the genus from southern 
Africa, but also the first record of the subfamily from the region. 

1 
Ann. S. Afr. Mm. 74 (1), 1977: 1-12, 6 figs. 



2 ANNALS OF THE SOUTH AFRICAN MUSEUM 

SYSTEMATIC PALAEONTOLOGY 

Class Cephalopoda Cuvier, 1797 

Order AMMONOIDEA Zittel, 1884 

Suborder Lytoceratina Hyatt, 1889 

Superfamily Tetragonitaceae Hyatt, 1900 

Family Tetragonitidae Hyatt, 1900 

Subfamily Gabbioceratinae Breistroffer, 1953 

The subfamily Gabbioceratinae has been the subject of two exhaustive 
accounts in recent years (Wiedmann 1962; Murphy 1967), and there is a diver- 
gence of opinion over which species should be referred to Gabbioceras versus 
Jauberticeras, as well as over the systematic position of a number of species 
regarded by Wiedmann as Gabbioceras and by Murphy as Eotetragonites. More 
serious is the disputation of the position of Parajaubertella Matsumoto, 1943, 
regarded by Wiedmann as a synonym of Gabbioceras and by Murphy as an 
independent homoeomorphous offshoot of the Eogaudryceras stock. There is, 
however, agreement that the subfamily has its origin in the group of Eogaudry- 
ceras {Eogaudryceras) numidum (Coquand). From this the line of descent runs 
to Gabbioceras, with G. lamberti (Breistroffer) as the earliest known member of 
that genus giving rise to Jauberticeras via J. jaubertianum during early late, or 
late early Aptian time. Jauberticeras is confined to the late Aptian and Albian; 
Gabbioceras extends to the Cenomanian. 

Genus Jauberticeras Jacob, 1907 
Type species 

Ammonites jaubertianus d'Orbigny (1850: 200, figs 9-10) by original designa- 
tion (see Howarth 1974 for clarification of the date and place of publication of 
Jauberticeras and of Ammonites jaubertianus as type species). 

Diagnosis 

Relatively small cadicones with a funnel-shaped umbilicus, no flank and a 
marked angulation demarcating umbilicus from the broad, flattened venter. 
Shell surface lirate with constrictions (sometimes only visible on the mould). 
Suture with three umbilical lobes. 

Discussion 

The authors take this opportunity to illustrate (Fig. 1A-G) a plaster cast 
of the lectotype of J. jaubertianum, the original of which is from the Aptian of 
Hyeges, Basses Alpes, France. When compared with Gabbioceras (a typical 
example of which is shown in Fig. 2A-G), the two genera are best distinguished 
on whorl section (following Murphy 1967: 604): in Gabbioceras the venter is 
strongly curved (Fig. 2A, C, D, G): in Jauberticeras it is flat (Fig. 1 A-B, D-F). 



CRETACEOUS FAUNAS FROM SOUTH AFRICA J 

The angulation is commonly sharper in Jauberticeras when compared with early 
Gabbioceras (Murphy 1967, fig. 1). 

Occurrence 

Late Aptian to mid Albian of southern France, Spain, the Caucasus, north 
Africa, Madagascar and Zululand. 



s 



■** 



^ 



/ 




B 




/ 
i 







' ~ : 





Fig. 1. Jauberticeras jaubertianum (d'Orbigny). The lectotype from the Aptian of Hyeges, 
Basses Alpes, France (Peticlerc Collection, Sorbonne, Paris). A-C x 2; D-G x 1. 



ANNALS OF THE SOUTH AFRICAN MUSEUM 





B 



** -** 











Fig. 2. Gabbioceras lamberti (Breistroffer). BMNH CI 0780, from the Aptian of Castellane, 

Basses Alpes, France. A-C x 2; D-G x 1. 



CRETACEOUS FAUNAS FROM SOUTH AFRICA J 

Jauberticeras collignoni Murphy, 1967 
Figs 3-5 

Jauberticeras latecarinatum Collignon non Anthula, J 963: 28, pi. 251 (fig. 1080). 
Jauberticeras collignoni Murphy, 1967: 606, pi. 64 (fig. 16). 

Halo type 

M. Collignon Collection no. 435, the original of Collignon (1963: 28, pi. 251 
(fig. 1080)) by monotypy. Reproduced here as Fig. 3A-H. From the Albian 
sensu lato of Betioky, Madagascar. 

Material 

Specimen D293 in the E. C. N. van Hoepen Collection. 

Emended diagnosis 

A Jauberticeras with a blunt lateral angle, moderately expanding whorls, 
weak lirae and irregularly spaced prorsiradiate convex constrictions and a 
prominent second saddle on the internal suture. 

Description 

Specimen D293 is a beautifully preserved mould of a phragmocone 21,5 mm 
in diameter, infilled with sparry calcite, and retaining a short portion of the body 
chamber filled with black quartzose glauconitic sandstone. Traces of the original 
shell material with delicate external ornament are preserved on the umbilical 
wall, but on the outer surface the shell is partially or wholly absent. Dimensions 
of the specimen, and others with which it is compared, are shown below. 
(D = diameter; Wb = whorl breadth; Wh = whorl height; U — umbilical 
diameter from seam to seam; K — umbilical diameter from keel to keel. 
Figures in parentheses are dimensions expressed as percentages of the total 
diameter.) 





D 


Wb 


Wh 


Wb:Wh 


U 


K 


J. collignoni 


22,4 (100) 


16,3 (74) 


6,9(31) 


2,36 


9,8 (44) 


18,7(84) 


D293 


18,2(100) 


13,3 (73) 


5,5 (30) 


2,41 


7,4 (41) 


14,2 (78) 


MC435 


19,3 (100) 


13,2(68) 


5,2 (27) 


2,54 


8,1 (42) 


14,9 (77) 



J. villoutreysi 

BMNHC68145 27,8(100) 18,5(67) 8,8(32) 2,13 12,0(43) 22,5(81) 

J. jaubertianum 

lectotype 32,0(100) 21,0(82) 10,5(32) 2,48 10,8(33) - (-) 

The coiling is very evolute, so that the angulation of the successive whorls 
is exposed in the umbilicus, giving it terraced, rather than evenly sloping sides. 
The umbilical wall is gently convex, whilst the angulation between it and the 
venter is abruptly rounded and blunt. The venter is broadly and evenly rounded, 
the distance between angulations comprising 84 per cent of the maximum 
diameter. The whorls expand quite slowly, so that there is a distinct tapering 



6 ANNALS OF THE SOUTH AFRICAN MUSEUM 

of the whorls towards the apex in both dorsal and ventral views. The mould 
bears low, faint, irregular folds and interspaces, which are markedly prorsiradiate 
on the flanks and sweep across the venter with a broad and shallow convex 
curvature. Two of these folds are strengthened into broad collars, one at the 
beginning of the body chamber, the other at a diameter of just over 20 mm. 
These are succeeded by distinct if shallow constrictions. 

On the venter, patches of partially exfoliated shell appear smooth, save for 
the low folds already mentioned; a few tiny areas preserve fine, dense lirae, 
invisible to the naked eye. In contrast, areas of umbilical wall where shell is 
well preserved show distinct if fine prorsiradiate lirae, concave close to the 
umbilical seam, but weakly flexed so as to be gently convex across the angulation. 
So far as can be discerned the whole of the shell was finely lirate, lirae paralleling 
the folds and constrictions visible on the mould. 

The external suture is well exposed (Fig. 4) and is deeply and intricately 
subdivided, as is characteristic of other species of the genus. The median saddle 
of the external lobe is prolonged into a series of narrow, tapering processes 
which strongly recall the siphonal band of certain tetragonitids (Kennedy & 
Klinger 1977). 






B 







H 




Fig. 3. Jauberticeras collignoni Murphy. Holotype, M. Collignon Collection no. 435, from the 
Albian sensu lato of Betioky, Madagascar. A-D x 2; E-H x 1. 



CRETACEOUS FAUNAS FROM SOUTH AFRICA 










m JBSC 



> * 1 '" > - *!■ . x l?' . * .. 



B 



Fig. 4. Sutures of Jauberticeras collignoni Murphy. Based on specimen D293 (Van Hoepen 
Collection) from the Mzinene Formation of the Mzinene River, Zululand, and of late middle 
or early late Albian age. A x 12; B x 6 to show details of siphonal structures. 



ANNALS OF THE SOUTH AFRICAN MUSEUM 






B 






"*>■ ■ <*",'" j<* 




H 



Fig. 5. Jauberticeras collignoni Murphy. D293 (Van Hoepen Collection) from the Mzinene 
Formation of the Mzinene River, Zululand, and of late middle or early late Albian age. 

A-D x 2; E-H x 1. 



CRETACEOUS FAUNAS FROM SOUTH AFRICA V 

Discussion 

Murphy based this species on the specimen figured by Collignon (1963, 
pi. 251 (fig. 1080)) which he refigured in dorsal view only (Murphy 1967, pi. 64 
(fig. 16), text-fig. 1 (9)). The original description is short, and in it Murphy 
characterized the species by its low expansion rate, smooth shell, and strong 
second saddle in the internal suture, lack of lirae and constrictions and similar 
whorl section to that of the type species. As can be seen from the holotype, here 
reillustrated (Fig. 3), the specimen does not quite fit Murphy's description in 
possessing constrictions. There are at least three distinct constrictions on the 
outer whorl, markedly prorsiradiate on the flank, and feebly convex across the 
venter, each being preceded by a slight swelling. The whorl section, with a 
breadth to height ratio of 2,54 at the maximum preserved diameter and 2,08 
360° before this, comprises only 68 per cent of the total diameter whereas in 
/. jaubertianum, to which it was said to be similar, the figure is 71 to 94 per cent. 
The whorl angulation is also significantly blunter. The latter, however, may be 
partially due to preservation or even of intra-specific value only, being dependent 
on ontogeny (see Murphy 1967, pi. 64 (fig. 23)). 

The second internal saddle is, however, prominently developed, whilst the 
shell surface is smooth, even under a lens. Close inspection shows that this may 
be no more than an artefact of preservation; the originally aragonitic shell has 
been replaced by iron-stained sparry calcite, and the whole of the surface of the 
umbilicus of the specimen is corroded and pitted. The venter is similarly pitted, 
and in places the angulation is covered and accentuated by traces of a succeeding 
whorl, now lost. This preservation and the remaining adherent matrix of fine 
sandstone suggest the specimen to be from a precisely similar setting to many 
of the Albian and Cenomanian ammonites from Zululand which the authors 
have studied, especially those found weathering out along the terraces of the 
Mzinene and on the Skoenberg, where identically preserved specimens of diverse 
ammonites have lost delicate growth lines and lirae by post-mortem processes. 
Lack of liration is not, therefore, a diagnostic feature of the species in the 
authors' view. With these qualifications in mind, the Zululand specimen, although 
larger than the Madagascan type, can be referred to Jauberticeras collignoni on 
the basis of similarities of whorl section, bluntness of angulation, relative propor- 
tions (which are similar but not identical), expansion rate, form and direction of 
constrictions, and associated collar-like swellings. Because the present specimen 
does not show the septal face (retaining as it does part of the body chamber) 
the nature of the internal suture is not determinable. 

The principal difference between J. collignoni and the type species, J. jauber- 
tianum (of which /. latecarinatum (Anthula) (1899: 101, pi. 7 (fig. 2a-c)) is 
probably a synonym according to Murphy (1967: 605-6)), lies in the more 
prominent development of the second saddle in the internal suture. 

Jauberticeras villoutreysi Murphy (1967: 606, pi. 64 (figs 25-29)) (the holo- 
type BMNH C68148 is reproduced here as Fig. 6A-G) has much coarser lirae 
on the umbilical wall, which give a distinctive appearance to the whorl angulation 



10 



ANNALS OF THE SOUTH AFRICAN MUSEUM 




f 




B 



•# 






iM 




Fig. 6. Jauberliceras villoutreysi Murphy. Holotype, BMNH C68I48, from the Albian of Clar, 
Alpes-Maritimes, France (Astier Collection). A-C x 2; D-G x 1. 



CRETACEOUS FAUNAS FROM SOUTH AFRICA 1 1 

(Fig. 6A). In addition, the expansion rate is such that the last half whorl is 
almost parallel-sided in profile (Fig. 6B, F), the venter flatter, and constrictions 
straight across the venter with a broad, band-like collar associated. In addition 
the saddles in the internal suture are of different sizes (Fig. 6C). 

Jauberticeras lantemoi Wiedmann (1962: 32, pi. 2 (fig. 2)) has quite different 
proportions and whorl section, and may possibly be a juvenile Gabbioceras 
(M. A. Murphy pers. comm. 7.3.1977). 

J. subbeticum Wiedmann (1962: 31, pi. 2 (fig. 3a-c)) has much coarser 
umbilical liration and a different form of whorl section and constrictions, and 
again is of dubious generic affinity. According to Murphy (pers. comm. 7.3. 1 977) 
it is probably a Scaphites. 

Occurrence 

The holotype of Jauberticeras collignoni is from the Albian sensu lato of 
Collignon's locality 435, 'Pied Est du Vohimaranitra (Betioky), Madagascar'. 
The Zululand specimen is from the Albian part of the Mzinene Formation along 
the Mzinene River near Hluhluwe. Precise locality data are lacking, but preserva- 
tion suggests it to be from locality 51 of Kennedy & Klinger (1974: 288, fig. 4) 
where concretions are exposed in the stream bed and bank extending around 
the broad meander of the river east-north-east of the Sisal Factory at 27°53'43"S, 
32°19'22"E. The specimen is thus of late middle or early late Albian age (Albian 
IV-V). 

ACKNOWLEDGEMENTS 

Our best thanks are due to General M. Collignon of Moirans (Jsere), who 
kindly allowed us to study the holotype of Jauberticeras collignoni, and has 
helped us throughout our work on the Zululand Cretaceous. We are also 
grateful to Dr M. K. Howarth and Mr D. Phillips (British Museum, Natural 
History) for allowing us to study the specimens in their care, and Dr J. Sornay 
of the Museum Nationale d'Histoire Naturelle, Paris, for help in our study of 
specimens in the D'Orbigny collection. 

REFERENCES 

Anthula, D. J. 1899. Ueber die Kreide Fossilien des Kaukasus. Beitr. Paldont. Geo/. Ost. Ung. 

12: 54-159. 
Collignon, M. 1963. Atlas des Fossiles caracteristiques de Madagascar {Ammonites), Fascicule 

X (Albien). Tananarive: Service Geologique. 
Howarth, M. K. 1974. The Lower Cretaceous ammonite genera proposed by C. Jacob in 

1907. Palaeontology 17: 727-728. 
Kennedy, W. J. & Klinger, H. C. 1974. Cretaceous faunas from Zululand and Natal, South 

Africa. Introduction, Stratigraphy. Bull. Br. Mus. nat. Hist. (Geol.) 25: 263-315. 
Kennedy, W. J. & Klinger, H. C. 1977. Cretaceous faunas from Zululand and Natal, South 

Africa. The ammonite family Tetragonitidae Hyatt, 1900. Ann. S. Afr. Mus. 73: 149-197. 
Kennedy, W. J. & Klinger, H. C. In press. Cretaceous faunas from Zululand and Natal, 

South Africa. The ammonite family Gaudryceratidae Spath, 1927. Bull. Br. Mus. nat. Hist. 

(Geol.). 



12 ANNALS OF THE SOUTH AFRICAN MUSEUM 

Murphy, M. A. 1967. The Ammonoid subfamily Gabbioceratinae Breistroffer. J. Paleont. 41: 

595-607. 
Orbigny, A. D\ 1850. Note sur quelques nouvellesespecesremarquablesdesetagesNeocomien 

et Aptien de France. J. Conchyliol. 1: 196-201. 
WitDMANN, J. 1962. Die Gabbioceratinae Breistroffer. NeuesJb. Geo/. Palaont. Abh. 115: 1-43. 



6. SYSTEMATIC papers must conform to the International code of zoological nomenclature 
(particularly Articles 22 and 51). 

Names of new taxa, combinations, synonyms, etc., when used for the first time, must be 
followed by the appropriate Latin (not English) abbreviation, e.g. gen. nov., sp. nov., comb, 
nov., syn. nov., etc. 

An author's name when cited must follow the name of the taxon without intervening 
punctuation and not be abbreviated; if the year is added, a comma must separate author's 
name and year. The author's name (and date, if cited) must be placed in parentheses if a 
species or subspecies is transferred from its original genus. The name of a subsequent user of 
a scientific name must be separated from the scientific name by a colon. 

Synonymy arrangement should be according to chronology of names, i.e. all published 
scientific names by which the species previously has been designated are listed in chronological 
order, with all references to that name following in chronological order, e.g. : 

Family Nuculanidae 

Nuculana (Lembulus) bicuspidata (Gould, 1845 

Figs 14-1 5 A 
Nucula (Leda) bicuspidata Gould, 1845: 37. 
Leda plicifera A. Adams, 1856: 50. 

Laeda bicuspidata Hanley, 1859: 118, pi. 228 (fig. 73). Sowerby, 1871 : pi. 2 (fig. 8a-b). 
Nucula largillierti Philippi, 1861: 87. 
Leda bicuspidata: Nickles, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9. 

Note punctuation in the above example : 

comma separates author's name and year 

semicolon separates more than one reference by the same author 

full stop separates references by different authors 

figures of plates are enclosed in parentheses to distinguish them from text-figures 

dash, not comma, separates consecutive numbers 

Synonymy arrangement according to chronology of bibliographic references, whereby 
the year is placed in front of each entry, and the synonym repeated in full for each entry, is 
not acceptable. 

In describing new species, one specimen must be designated as the holotype; other speci- 
mens mentioned in the original description are to be designated paratypes ; additional material 
not regarded as paratypes should be listed separately. The complete data (registration number, 
depository, description of specimen, locality, collector, date) of the holotype and paratypes 
must be recorded, e.g. : 

Holotype 

SAM-A13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King's Beach 
Port Elizabeth (33°51'S 25°39'E), collected by A. Smith, 15 January 1973. 

Note standard form of writing South African Museum registration numbers and date. 

7. SPECIAL HOUSE RULES 

Capital initial letters 

(a) The Figures, Maps and Tables of the paper when referred to n the text 

e.g. '. . . the Figure depicting C. namacolus . . .'; '. . . in C. namacolus (Fig. 10) . . .' 

(b) The prefixes of prefixed surnames in all languages, when used in the text, if not preceded 
by initials or full names 

e.g. Du Toit but A. L. du Toit; Von Huene but F. von Huene 

(c) Scientific names, but not their vernacular derivatives 
e.g. Therocephalia, but therocephalian 

Punctuation should be loose, omitting all not strictly necessary 

Reference to the author should be expressed in the third person 

Roman numerals should be converted to arabic, except when forming part of the title of a 

book or article, such as 

'Revision of the Crustacea. Part VIII. The Amphipoda.' 
Specific name must not stand alone, but be preceded by the generic name or its abbreviation 

to initial capital letter, provided the same generic name is used consecutively. 
Name of new genus or species is not to be included in the title: it should be included in the 

abstract, counter to Recommendation 23 of the Code, to meet the requirements of 

Biological Abstracts. 



WILLIAM JAMES KENNEDY 

& 
HERBERT CHRISTIAN KLINGER 

CRETACEOUS FAUNAS FROM ZULULAND 

AND NATAL, SOUTH AFRICA 
A JAUBERTICERAS FROM THE MZINENE 

FORMATION (ALBIAN) 



J 



VOLUME 74 PART 2 



DECEMBER 1977 



ISSN 0303-2515 



MUS. COMP. ZOOU 



RY 



NOV J 9 iq fin 



H ., ^ 



ANNALS 




INSTRUCTIONS TO AUTHORS 

1. MATERIAL should be original and not published elsewhere, in whole or in part. 

2. LAYOUT should be as follows: 

(a) Centred masthead to consist of 

Title: informative but concise, without abbreviations and not including the names of new genera or species 

Author's(s') name(s) 

Address(es) of author(s) (institution where work was carried out) 

Number of illustrations (figures, enumerated maps and tables, in this order) 

(b) Abstract of not more than 200 words, intelligible to the reader without reference to the text 

(c) Table of contents giving hierarchy of headings and subheadings 

(d) Introduction 

(e) Subject-matter of the paper, divided into sections to correspond with those given in table of contents 

(f ) Summary, if paper is lengthy 

(g) Acknowledgements 
(h) References 

(i) Abbreviations, where these are numerous 

3. MANUSCRIPT, to be submitted in triplicate, should be typewritten and neat, double spaced 
with 2,5 cm margins all round. First lines of paragraphs should be indented. Tables and a list of 
legends for illustrations should be typed separately, their positions indicated in the text. All 
pages should be numbered consecutively. 

Major headings of the paper are centred capitals; first subheadings are shouldered small 
capitals ; second subheadings are shouldered italics ; third subheadings are indented, shouldered 
italics. Further subdivisions should be avoided, as also enumeration (never roman numerals) 
of headings and abbreviations. 

Footnotes should be avoided unless they are short and essential. 

Only generic and specific names should be underlined to indicate italics; all other marking 
up should be left to editor and publisher. 

4. ILLUSTRATIONS should be reducible to a size not exceeding 12x18 cm (19 cm including 
legend); the reduction or enlargement required should be indicated; originals larger than 
35 x 47 cm should not be submitted; photographs should be rectangular in shape and final 
size. A metric scale should appear with all illustrations, otherwise magnification or reduction 
should be given in the legend ; if the latter, then the final reduction or enlargement should be 
taken into consideration. 

All illustrations, whether line drawings or photographs, should be termed figures (plates 
are not printed; half-tones will appear in their proper place in the text) and numbered in a 
single series. Items of composite figures should be designated by capital letters; lettering of 
figures is not set in type and should be in lower-case letters. 

The number of the figure should be lightly marked in pencil on the back of each illustration. 

5. REFERENCES cited in text and synonymies should all be included in the list at the end of 
the paper, using the Harvard System (ibid., idem, he. cit., op. cit. are not acceptable): 

(a) Author's name and year of publication given in text, e.g. : 

'Smith (1969) describes . . .' 

'Smith (1969: 36, fig. 16) describes 

'As described (Smith 1969a, 19696; Jones 1971)' 

'As described (Haughton & Broom 1927) ..." 

'As described (Haughton et al. 1927) . . .' 

Note: no comma separating name and year 

pagination indicated by colon, not p. 

names of joint authors connected by ampersand 

et al. in text for more than two joint authors, but names of all authors given in list of references. 

(b) Full references at the end of the paper, arranged alphabetically by names, chronologically 
within each name, with suffixes a, b, etc. to the year for more than one paper by the same 
author in that year, e.g. Smith (1969a, 19696) and not Smith (1969, 1969a). 

For books give title in italics, edition, volume number, place of publication, publisher. 

For journal article give title of article, title of journal in italics (abbreviated according to the World list o y 
scientific periodicals. 4th ed. London: Butterworths, 1963), series in parentheses, volume number, part 
number (only if independently paged) in parentheses, pagination (first and last pages of article). 

Examples (note capitalization and punctuation) 

Bullough, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan. 

Fischer, P.-H. 1948. Donnees sur la resistance et de le vitalite des mollusques. /. Conch., Paris 88: 100-140. 

Fischer, P.-H., Duval, M. & Raffy, A. 1933. Etudes sur les echanges respiratoires des littorines. Archs 

Zool. exp. gen. 74: 627-634. 
Kohn, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon. 

Ann. Mag. not. Hist. (13) 2: 309-320. 
Kohn, A. J. 19606. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean. 

Bull. Bingham oceanogr. Coll. 17 (4): 1-51. 
Thtele, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. In: Schultze, L. Zoologische 

und anthropologische Ergebnisse einer Forschungsreise im west lichen und zentralen Sud-Afrika 4: 269-270. 

Jena: Fischer. Denkschr. med.-naturw. Ges. Jena 16: 269-270. 

(continued inside back cover) 



ANNALS OF THE SOUTH AFRICAN MUSEUM 
ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM 



Volume 74 Band 
December 1977 Desember 
Part 2 Deel 




THE SOUTH AFRICAN MUSEUM'S 

MEIRING NAUDE CRUISES 

PART 5 

CRUSTACEA, DECAPODA, REPTANTIA 

AND NATANTIA 



By 



BRIAN KENSLEY 



Cape Town Kaapstad 



The ANNALS OF THE SOUTH AFRICAN MUSEUM 

are issued in parts at irregular intervals as material 
becomes available 

Obtainable from the South African Museum, P.O. Box 61, Cape Town 

Die ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM 

word uitgegee in dele op ongereelde tye na beskikbaarheid 

van stof 

Verkrygbaar van die Suid-Afrikaanse Museum, Posbus 61, Kaapstad 



OUT OF PRINT/UIT DRUK 

1, 2(1-3, 5-8), 3(1-2, 4-5, 8, t.-p.i.), 5(1-3, 5, 7-9), 

6(1, t.-p.i.), 7(1-4), 8, 9(1-2, 7), 10(1-3), 
11(1-2, 5, 7, t.-p.i.), 15(4-5), 24(2), 27, 31(1-3), 33 



Price of this part/Pry s van hierdie deel 
R3,00 



Trustees of the South African Museum © Trustees van die Suid-Afrikaanse Museum 

1977 



ISBN 908407 27 



Printed in South Africa by In Suid-Afrika gedruk deur 

The Rustica Press, Pty., Ltd., Die Rustica-pers, Edms., Bpk., 

Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap 



THE SOUTH AFRICAN MUSEUM'S MEIRING NAUDE CRUISES 

PART 5 
CRUSTACEA, DECAPODA, REPTANTIA AND NATANTIA 

By 

Brian Kensley 

South African Museum, Cape Town 

(With 16 figures) 
[MS. accepted 10 May 1977] 

ABSTRACT 

Sixty-one species of Reptantian and Natantian decapod crustaceans from deep water off 
Natal are listed. Of the thirteen new records of Natantia dealt with from the area, three are 
described as new species. These are Benthesicymus expansus, Hymenopenaeus kannemeyeri and 
Pasiphaea meiringnaudei. 



CONTENTS 






PAGE 


Introduction 


. 13 


Species List 


. 13 


Systematic Discussion . 


. 20 


Acknowledgements 


. 43 


References . 


. 43 



INTRODUCTION 

The present paper forms part of a series based on material collected by the 
South African Museum during two cruises off the Natal coast, on the R/V Meiring 
Naude (see Fig. 1). For the scope of, and the background to, this programme, 
as well as for all station data, the reader is referred to Louw (1977). 

All the species of Reptantia and Natantia collected are listed systematically, 
but only new species, new records for South Africa, and a few additional species 
requiring further comment are dealt with in the systematic text. 

The following abbreviations are used throughout this paper: SAM— South 
African Museum catalogue number; SM— Meiring Naude station numbers; 
CB— carapace breadth; CL— carapace length; RL— rostral length; TL— total 
length ; ovig. — ovigerous ; juv. —juvenile. 

SPECIES LIST 

Where specimens are not identified to species level, these were either immature or damaged. 
' * New record for South African waters. 

SM Station ovig. 



PALINURA 

Family Eryonidae 

Stereomastis sculpt a (Smith) . . . . 58 
(Barnard 1950: 572) 83 

{Eryoneicus caecus) . . . .107 

13 
Ann. S. Afr. Mus. 74 (2), 1977: 13-44, 16 figs. 



no. <$$ ?$ $? juv. 



14 



ANNALS OF THE SOUTH AFRICAN MUSEUM 



3T 10' 20' 30' 40' 50' 32° 10' 20' 30' W 50' 33° 10' 20' 




50' 31° Iff 20' 30' 40' 50' 32° 10' 20' 30' 40' 50' 330 10' 20 



Fig. 1. Map showing localities of collecting stations. 



THE SOUTH AFRICAN MUSEUM'S MEIRING NAUDE CRUISES 



15 



SM Station 

no. (JcJ 



9? 



ovig. 

?¥ 



Family Palinuridae 

Palinurus delagoae Barnard . 

(Berry & Plante 1973: 374) 
Projasus parked (Stebbing) . 

(George & Grindley 1964: 87) 

ASTACURA 

Family Astacidae 
Nephropsis atlantica Norman . 
(Barnard 1950: 530) 

NATANTIA PENAEIDEA 

Family Penaeidae 

Subfamily Sicyoninae 
Sicyonia longicauda Rathbun . 
(Barnard 1950: 635) 

Subfamily Aristeinae 
Aristaeomorpha foliacea (Risso) 
(Barnard 1950: 625) 



juv. 



*Aristeus semidentatus (Bate) .... 

(see systematic section, p. 20) 
*Aristeus virilis (Bate) ..... 

(see systematic section, p. 21) 

* Benthesicymus expansus sp. nov. . 

(see systematic section, p. 22) 
* Benthesicymus investigatoris Alcock & Anderson 

(see systematic section, p. 26) 



Gennadas bouvieri Kemp 
(Kensley 19716: 273) 



Gennadas capensis Caiman 
(Kensley 19716: 277) 

Gennadas clavicarpus De Man 
(Kensley 19716: 278) 



Gennadas gilchristi Caiman 
(Kensley 19716: 280) 



22 
38 



38 
66 



86 



I 



7 


1 


2 


22 


2 


4 


28 


1 


— 


38 


3 


3 


44 


— 


2 


66 


— 


1 


83 


1 


— 


92 


— 


2 


99 


1 


— 


100 


3 


3 


107 


12 


— 


58 


2 





91 


1 


— 


107 


— 


1 


107 


1 


— 


38 





1 


58 


— 


1 


66 


— 


1 


107 


— 


2 


55 


— 


1 


56 


1 


— 


63 


— 


4 


88 


— 


1 


105 


— 


1 


111 


— 


1 


99 


— 


1 


105 


1 


1 


55 


1 


1 


63 


— 


1 


88 


— 


1 


96 


1 


1 


105 


— 


1 


111 


3 


— 


25 


— 


3 


33 


2 


4 


49 


— 


2 


55 


3 


4 



16 



ANNALS OF THE SOUTH AFRICAN MUSEUM 



Gennadas parvus Bate . 
(Kensley 19716: 287) 



Gennadas scutatus Bouvier 
(Kensley 19716: 288) 



Gennadas tinayrei Bouvier 
(Kensley 19716: 290) 



Plesiopenaeus edwardsianus (Johnson) 
(Crosnier & Forest 1973: 292) 



Plesiopenaeus niiidus Barnard 
(Barnard 1950: 622) 



Subfamily Solenocerinae 
Haliporoides triarthrus (Stebbing) 

(Barnard 1950: 619) 
* Hymenopenaeus kannemeyeri sp. nov. 

(see systematic section, p. 27) 
Hymenopenaeus sp. 



SM Station 






ovig. 


no. 


(W 


?? 


?? 


56 


15 


12 


— 


63 


— 


6 


— 


88 


9 


4 


— 


96 


— 


3 


— 


97 


5 


3 


— 


105 


11 


8 


— 


111 


3 


1 


— 


112 


7 


3 


— 


25 


1 


— 


— 


56 


2 


— 


— 


63 


5 


3 


— 


105 


— 


1 


— 


111 


1 


1 


— 


112 


1 


— 


— 


25 


1 


— 


— 


33 


— 


1 


— 


56 


1 


— 


— 


63 


— 


2 


— 


88 


— 


1 


— 


97 


1 


— 


— 


105 


— 


1 


— 


112 


3 


- 


— 


56 


1 


1 


— 


71 


— 


1 


— 


88 


2 


4 


— 


96 


2 


1 


— 


105 


— 


2 


— 


111 


1 


— 


— 


112 


1 


1 


— 


56 


1 


1 


— 


63 


3 


— 


— 


88 


5 


— 


— 


96 


2 


1 


— 


7 


— 


1 


— 


28 


1 


— 


— 


38 


— 


3 


— 


58 


1 


1 


— 


74 


2 


— 


— 


107 


1 


— 


— 


7 


3 


4 


— 


22 


1 


7 


— 


28 


1 


1 


— 


31 


— 


1 


— 


38 


3 


5 


— 


58 


3 


7 


— 


66 


2 


2 


— 


83 


1 


7 


— 


92 


— 


2 


— 


99 


— 


1 


— 


107 


7 


34 


— 


22 


5 


3 


_ 


92 


1 


— 


— 


38 


1 


1 


— 



JUV. 






THE SOUTH AFRICAN MUSEUM'S MEIR1NG NAUDE CRUISES 



17 



Subfamily Penaeinae 
*Funchalia villosa (Bouvier) . 
(see systematic section, p. 29) 



Parapenaeopsis acclivirostris Alcock 

(Barnard 1950: 604) 
Penaeopsis rectacuta (Bate) . 

(Hall 1962: 18) 
Penaeus indicus Edwards 

(Barnard 1950: 588) 

Family Sergestidae 

Sergestes {Sergestes) armatus Kroyer 
(Kensley 1971a: 232) 



Sergestes {Sergestes) atlanticus Edwards 
(Kensley 1971a: 234) 



Sergestes {Sergestes) curvatus Crosnier & Forest 
(Crosnier & Forest 1973: 315) 



Sergestes {Sergestes) orientalis Hansen 
(Kensley 1971a: 238) 



Sergestes {Sergestes) sargassi Ortmann 
(Kensley 1971a: 241) 



Sergestes {Sergestes) pectinatus Sund 
(Kensley 1971a: 240) 



SM Station 






ovig, 


no. 


c?c? 


99 


99 


18 


— 


1 


— 


33 


— 


1 


— 


88 


1 


1 


— 


111 


1 


— 


— 


115 


— 


2 


— 


15 


3 


11 


— 


114 


1 


— 


— 


18 




1 




33 


2 


2 


— 


49 


1 


1 


— 


55 


1 


2 


— 


56 


— 


1 


— 


63 


2 


— 


— 


88 


2 


3 


— 


96 


— 


2 


— 


105 


— 


1 


— 


56 


— 


1 


— 


63 


1 


11 


— 


88 


2 


3 


— 


105 


1 


1 


— 


111 


1 


— 


— 


112 


3 


1 


— 


>t 5 


— 


1 


— 


18 


— 


2 


— 


25 


3 


— 


— 


33 


1 


1 


— 


49 


1 


1 


— 


55 


1 


— 


— 


56 


1 


2 


— 


63 


2 


— 


— 


88 


1 


— 


— 


97 


— 


1 


— 


111 


2 


1 


— 


112 


1 


2 


— 


55 


— 


3 


— 


88 


— 


3 


— 


96 


1 


2 


— 


97 


1 


1 


— 


105 


1 


— 


— 


112 


3 


1 


— 


25 


— 


1 


— 


56 


— 


1 


— 


63 


1 


— 


— 


88 


1 


— 


— 


105 


1 


— 


— 


111 


— 


2 


— 


112 


1 


— 


— 


88 


1 


1 


— 


96 


2 


— 


— 


111 


— 


2 


— 


112 


1 


1 


— 



JUV. 



18 



ANNALS OF THE SOUTH AFRICAN MUSEUM 





SM Station 






ovig 




no. 


SS 


?? 


?? 


Sergestes (Sergio) laminatus Burkenroad . 


56 


2 


— 


— 


(Kensley 1971a: 251) 


88 


3 


4 


— 




97 


4 


1 


— 




105 


5 


5 


— 




111 


1 


2 


— 




112 


1 


3 


— 


Sergestes (Sergio) potens Burkenroad 


7 


— 


2 


— 


(Kensley 1971a: 253) 










Sergestes (Sergio) prehensilis Bate . 


5 


— 


2 


— 


(Kensley 1971a: 253) 


18 


— 


2 


— 




25 


13 


19 


— 




33 


2 


5 


— 




49 


1 


— 


— 




55 


1 


1 


— 




56 


3 


— 


— 




63 


5 


1 


— 




67 


1 


2 


— 




88 


4 


4 


— 




96 


7 


2 


— 




97 


2 


2 


— 




105 


3 


3 


— 




111 


1 


1 


— 




112 


7 


3 


— 


Sergestes (Sergio) regalis Gordon . 


25 


— 


1 


— 


(Kensley 1971a: 256) 


55 


— 


1 


— 




56 


1 


— 


— 




88 


— 


1 


— 




96 


1 


— 


— 




99 


— 


1 


— 




112 


— 


1 


— 


Sergestes (Sergio) scintillans Burkenroad . 


25 


1 


— 


— 


(Kensley 1971a: 257) 


88 


1 


2 


— 




96 


1 


1 


— 




105 


3 


— 


— 




112 


2 


4 


— 


Sergestes (Sergio) splendens Sund . 


88 


2 


— 


— 


(Kensley 1971a: 260) 


96 


— 


2 


— 


* Sergestes (Sergio) talismani Barnard 


55 


2 


2 


— 


(see systematic section, p. 31) 


63 


— 


1 


— 




88 


4 


— 


— 




97 


— 


1 


— 




112 


— 


3 


— 


NATANTIA CARIDEA 










Family Oplophoridae 










* Acanthephyra armata Edwards 


38 


4 


4 


— 


(see systematic section, p. 31) 


58 


3 


3 


— 


Acanthephyra eximia Smith . 


3 


— 


1 


— 


(Crosnier & Forest 1973: 34) 


38 


— 


8 


— 




44 


— 


1 


— 




58 


3 


5 


— 




99 


— 


1 


— 




107 


1 


1 


— 


* Acanthephyra indica Balss 


56 


1 


— 


— 


(see systematic section, p. 31) 


88 


— 


— 


1 




96 


— 


1 


— 




105 


2 


— 


— 



JUV. 






THE SOUTH AFRICAN MUSEUM'S MEIRING NAUDE CRUISES 



19 



Acanthephyra quadrispinosa Kemp 
(Barnard 1950: 668) 



Oplophorus gracilirostris Edwards 

(Chace 1936: 30) 
Oplophorus spinicauda Edwards 

(Kensley 1969: 169) 



Oplophorus spinosus (Brulle) . 

(Crosnier & Forest 1973: 25) 
Systellaspis debilis (Edwards) 

(Crosnier & Forest 1973: 87) 



Family Nematocarcinidae 
Nematocarcinus sp. 



Family Pasiphaeidae 

*Eupasiphae gilesii Wood-Mason 
(see systematic section, p. 32) 

*Pasiphaea sivado (Risso) , . 
(see systematic section, p. 32) 

*Pasiphaea meiringnaudei sp. nov. 
(see systematic section, p. 34) 



Family Hippolytidae 

Hippolysmata tugelae Stebbing 
(Barnard 1950: 712) 



SM Station 






ovig. 




no. 


S3 


?? 


9? 


juv 


Ill 


— 


— 


1 


— 


112 


— 


— 


1 


— 


5 


— 


1 


1 


— 


18 


2 


2 


2 


— 


25 


1 


3 


18 


— 


33 


3 


7 


7 


— 


49 


3 ' 


5 


5 


— 


55 


5 ■ 


1 


— 


9 


56 


7 


— 


5 


20 


63 


1 


7 


10 


10 


88 


3 


4 


2 


14 


96 


3 


— 


1 


2 


97 


3 


1 


1 


4 


105 


4 


1 


— 


8 


111 


7 


1 


— 


7 


112 


3 


— 


2 


19 


33 


1 


— 


— 


— 


63 


1 


— 


— 


— 


18 


— 


1 


— 


3 


25 


— 


2 


— 


— 


56 


— 


— 


1 


— 


111 


— 


2 


— 


1 


112 


— 


3 


— 


1 


33 


— 


2 


— 


1 


25 


— 


— 


— 


1 


5 


1 


— 


4 


— 


18 


4 


2 


1 


— 


25 


13 


10 


14 


— 


33 


7 


3 


3 


— 


49 


5 


4 


1 


— 


55 


3 


— 


1 


— 


56 


— 


1 


1 


— 


63 


— 


1 


— 


— 


88 


— 


— 


— 


1 


105 


— 


— 


— 


1 


111 


2 


— 


1 


— 


112 


1 


— 


1 


5 


7 


1 











38 


1 


— 


— 


— 


66 


— 


— 


1 


— 


90 


1 


— 


— 


— 


92 


— 


— 


1 


— 


105 


1 


— 


— 


— • 


63 





2 








85 


— 


1 


— 


— 


28 


— 


1 


— 


— 


38 


1 


— 


— 


— 


40 


— 


2 


— 


— 


58 


4 


— 


— 


— 


91 


— 


1 


— 


— 


107 


— 


— 


5 


— 



115 



20 



ANNALS OF THE SOUTH AFRICAN MUSEUM 



Family Pandalidae 
Heterocarpus dorsalis Bate 
(see systematic section, p. 



38) 



Heterocarpus laevigatus Bate . 
(see systematic section, p. 38) 

Heterocarpus tricarinatus Alcock & Anderson 

(see systematic section, p. 38) 
Parapandalus richardi (Couti6re) . 



Plesionika martia (Edwards) . 

(Crosnier & Forest 1973: 212) 



Family Glyphocrangonidae 
Glyphocrangon dentatus Barnard 
(see systematic section, p. 40) 



* Glyphocrangon regalis Bate . 
(see systematic section, p. 40) 



Family Crangonidae 
Pontophilus sculptus (Bell) 
(Barnard 1950: 810) 



SM Station 






ovig. 


no. 


<?<? 


?? 


?? 


7 


1 


— 


— 


28 


— 


1 


— 


38 


2 


1 


— 


75 


1 


— 


— 


83 


3 


— 


— 


38 


— 


1 


— 


66 


1 


— 


— 


99 


1 


1 


1 


22 


— 


1 


1 


56 


1 








63 


1 


— 


— 


88 


1 


— 


— 


111 


1 


— 


1 


7 


1 


1 


— 


22 


— 


— 


2 


38 


2 


1 


2 


58 


— 


— 


1 


66 


2 


1 


2 


83 


— 


— 


2 


100 


— 


— 


1 


22 






8 


28 




— 


— 


38 




— 


— 


66 




1 


— 


100 




— 


— 


7 




1 


1 


28 


— 


— 


1 


38 




2 


4 


44 




— 


— 


58 


— 


1 


— 


74 




— 


— 


83 




3 


1 


92 


— 


— 


2 


99 


— 


1 


— 


86 





1 






JUV. 












SYSTEMATIC DISCUSSION 

(new species, new records, and selected other species) 

PENAEIDEA 

Family Penaeidae 

Aristeus semidentatus (Bate) 

Figs 2A, 3A 

Hemipenaeus semidentatus Bate, 1888: 305, pi. 49 (fig. 1). 

Aristaeus semidentatus: Wood-Mason & Alcock, 1891: 280. Alcock, 1901: 31. Alcock & 

McArdle, 1901: pi. .49 (fig. 3). 
Aristeus semidentatus: De Man, 1911 : 29. 



THE SOUTH AFRICAN MUSEUM'S MEIRING NAUDE CRUISES 



21 



Previous records 

Laccadive Islands ; Cape Comorin ; Bay of Bengal ; off Banda ; Kei Islands ; 
Kermadec Islands. 



Material 

SAM-A15139 



SM 107 12 $$ CL + RL 26,8 mm x TL 77,0 mm to 
CL + RL 31,5 mm x 92,0 mm 



Remarks 

The genus Aristeus is characterized by the lack of an hepatic spine, an 
indistinct cervical groove, the third pereiopods possessing an epipodite but no 
podobranch, the fourth pereiopods lacking an epipodite, the exopodite of the 
second maxilliped longer and stronger than the exopodite of the third maxilliped. 

There is some variation in the rostral dentition of the present material, 
several specimens showing a small dorsal and ventral tooth close to the apex 
(Fig. 2 A). (Burucovsky & Romensky 1972, also describe rostral variability in 
Aristeus varidens.) 

Aristeus virilis (Bate) 
Figs 2B, 3B 

Hemipenaeus virilis Bate, 1888: 303, pi. 44 (fig. 4). 
Hemipenaeus tomentosus Bate, 1888: 307, pi. 49 (figs 2-3). 
Aristaeus virilis: Wood-Mason & Alcock, 1891: 279. Alcock, 1901: 30. 
Aristeus virilis: De Man, 1911 : 27. 





Fig. 2. A. Aristeus semidentatus. B. Aristeus virilis. 



22 



ANNALS OF THE SOUTH AFRICAN MUSEUM 



Previous records 

Andaman Sea; China Sea; Philippines; Bali Sea; New Hebrides. 



Material 

SAM-A15159 

SAM-A15160 

SAM-A15161 



SM 58 2 tfS CL 41,2 mm x TL 110,5 mm; 37,0 mm x 

± 95 mm (telson damaged) 

SM 91 1 S 43,0 mm x 109,0 mm 

SM 107 1 $ (rostrum damaged) 



Remarks 

The present material shows the characteristic thickening of the distal part 
of the scaphocerite, but the males lack the pubescence of the integument. A twist 
in the base of the inner antennular flagellum is also present. The two species of 
Aristeus dealt with here may be distinguished by the differences in the petasmata 
(Fig. 3) and rostra (Fig. 2), and the pubescent carapace of some specimens of 
A. virilis. 

Benthesicymus expansus sp. no v. 

Figs 4-5 
Description 

Male 

Carapace (Fig. 4A) integument soft, membranous. Rostrum an entire crest, 
apex missing in holotype, ventral border bearing slight ridge carrying setae. No 





Fig. 3. A. Aristeus semidentatus petasma. B. Aristeus virilis petasma. 



THE SOUTH AFRICAN MUSEUMS MEIRING NAUDE CRUISES 23 

antennal spine; orbitoantennal groove extended posteroventrally to branchial 
region; rounded ridge situated above orbitoantennal groove; branchiostegal 
spine small, supported by short carina, no hepatic spine; cervical groove strong, 
almost reaching orbitoantennal groove; suprabranchial ridge fairly strong, 
rounded. Thoracic sternite between fourth pereiopods bearing short narrowly 
rounded, forwardly directed lobe. Posterior half of third pleon segment, and 
pleon segments four to six with mediodorsal carina, none bearing posterior spine. 
Telson carrying four pairs of lateral spines on distal third, apex acute (Fig. 4B). 

Antennal scaphocerite (Fig. 5F) blade two and one-third times longer than 
wide, outer margin reinforced by tapering ridge, ending in small spine some 
distance from apex of scale; basal segment carrying small dorsal hook-like 
process. 

Antennular peduncle segments setose. 

Ocular tubercle present on inner margin of proximal half of eye-stalk. 

Mandibular palp two-segmented, basal segment one and a half times longer 
and considerably broader than distal segment; mandible consisting of strong, 
straight cutting edge and three small teeth on inner face (Fig. 5A). 

First maxilliped (Fig. 5B) exopod narrowing abruptly to segmented distal 
portion. Second maxilliped (Fig. 5E) with merus very expanded, distally rounded, 
inner margin setose. 

Third maxilliped longer than first pereiopod (Fig. 4A), merus and ischium 
expanded, fringed with setae on ventral margin ; propodus and carpus relatively 
slender, of equal length, dactylus broadest at midpoint, tapering distally. First 
pereiopod chela relatively slender, finger and thumb slightly longer than palm ; 
carpus slender; merus and ischium expanded. 

Second pereiopod (Fig. 4A) longer than first, very slender, finger and thumb 
of chela equal in length to palm; carpus and merus of equal length. 

Third to fifth pereiopods missing in holotype. 

Petasma slender (Fig. 4C), divided distally into median narrow lobe con- 
nected to truncate, rounded portion, and outer smoothly convex lobe bearing 
spinules. 

Inner scale of appendix masculina narrow, elongate, apically rounded, 
bearing short setae; outer scale broad, longer than inner. 

Material 

Holotype SAM-A15140 SM 107 1 $ CL 50,5 mm TL 150 mm 

Remarks 

The lack of mediodorsal pleonal spines, the ocular tubercle being proximal 
in position, the lack of an hepatic spine, and the presence of a mediodorsal keel 
on pleonal segments 3-6, place this specimen close to B. iridescens Bate (see 
Burkenroad 1936: 45; Roberts & Pequegnat 1970: 42). The latter authors note 
the difficulty of sorting out the 'Brasiliensis' group of species of Benthesicymus, 
due mainly to a lack of information on intraspecific variation. In spite of this 



24 



ANNALS OF THE SOUTH AFRICAN MUSEUM 







Fig. 4. Benthesicymus expansus sp. nov. A. Holotype, carapace in lateral view. B. Telson. 
C. Anterior and posterior views of petasma. 



THE SOUTH AFRICAN MUSEUM'S MEIRING NAUDE CRUISES 



25 



limitation, the present new species is described, albeit from a single specimen. 
The species has two features which easily separate it from B. iridescens. These 
are the non-dentate rostrum, and the expanded meri and ischia of the third 
maxilliped and first pereiopods. (The specific name is derived from this latter 
feature.) In addition, differences from B. iridescens are seen in the petasma, the 
appendix masculina, the telson, the lack of an antennal spine, and in the carapace 
sculpture. 









Fig. 5. Benthesicymus expansus sp. nov. A. Mandible. B. First maxilla. C. Second maxilla. 
D. First maxilliped. E. Second maxilliped. F. Antennal scaphocerite. 



26 ANNALS OF THE SOUTH AFRICAN MUSEUM 

Benthesicymus investigatoris Alcock & Anderson 

Fig. 6 

Benthesicymus investigatoris Alcock & Anderson, 1899: 282. Alcock, 1899: pi. 41 (fig. 2); 
1901 : 44. Burkenroad, 1936: 25. 

Previous records 

Saya de Malha (mid Indian Ocean); Andaman Sea; Gulf of Manar; 
Halmahera Sea; Ceram Sea; Hawaii. 

Material 

5 $$ CL + RL 24,0 mm x TL 76 to 31,0 mm x 89,5 mm 




Fig. 6. Benthesicymus investigatoris. A. Carapace in lateral view. B. Thelycum. 



THE SOUTH AFRICAN MUSEUM'S ME/R/NG NAVDF. CRUISES 27 

Hymenopenaeus kannemeyeri sp. no v. 

Fig. 7 
Description 

Carapace (Fig. 7A) firm, glabrous. Rostrum about one-third length of 
carapace directed obliquely upward at angle of about 1 5°, with eight dorsal 
teeth (three postorbital) in uninterrupted series, apical tooth very small, ventral 
margin bearing fringe of setae. Carapace with strong antennal and postantennal 
spine, each supported by stout carina; hepatic spine lacking keel; branchiostegal 
spine marginal, supported by keel extending posteriorly to hind margin of cara- 
pace; cervical sulcus well developed, with posterior margin of groove at midpoint 
marked by a short keel, joining the hepatic groove ventrally. Hepatic groove 
marked posteroventrally by short keel. Branchiocardiac sulcus well developed 
with supra-branchial portion having a straight, rounded ridge ventrally. First 
pleon segment dorsally rounded, segment two with posterior half keeled, third 
to sixth segments fully keeled, segments four to six with short posterior spine. 
Telson (Fig. 7D) with two fixed spines subterminally, with strong dorsolateral keel 
extending from spines to proximal margin; outer uropodal ramus just reaching 
telsonic apex. 

Eye reaching slightly beyond midpoint of rostrum, tubercle at midpoint of 
medial eye-stalk margin (Fig. 7E). 

Prosartema of antennule short and rounded, carrying long setae; basal 
peduncular segment with strong distal spine ; stylocerite a strong spine ; terminal 
peduncular segment with lateral setose ridge and thick setose patch dorsally; 
flagellum circular in cross-section. 

Scaphocerite extending well beyond antennular peduncle. 

Mandibular palp conspicuous, setose; distal segment slender, tapering, 
shorter than proximal segment, latter broadly oval. 

Third maxilliped extending slightly beyond scaphocerite. 

First pereiopod with strong distoventral spine on ischium and basis. 

Rudimentary exopods on first to fifth pereiopods and third maxilliped; 
epipods on second and third maxillipeds and first to fourth pereiopods. 

Inner margin of fifth pereiopod basis in male with anteriorly directed 
triangular lobe above genital aperture. 

Thelycum of female (Fig. 7C) relatively simple, sternites of fourth and fifth 
pereiopods with rounded, convex central areas. 

Petasma divided distally into two lobes; inner lobe longer than outer, 
truncate, with distal hollowed portion produced laterally into triangular point; 
outer lobe also truncate, with posterior strengthening ridge. Appendix masculina 
of second pleopod (Fig. 7F) about half length of appendix interna; both bearing 
marginal spinules; leaf-like expansion present on outer side of base of endopod. 

Material 

Holotype SAM-A15141 SM 38 $ CL + RL 43,5 mm TL 119,5 mm 
Allotype SAM-A15141 SM 38 ? CL + RL 39,8 mm TL 1 10,8 mm. 



28 



ANNALS OF THE SOUTH AFRICAN MUSEUM 



Remarks 

The present specimens with the rostral and post-rostral teeth not separated, 
with a strong branchiostegal spine, and lacking a pterygostomian spine, fall into 
Burkenroad's (1936) Group II. 

This group includes H. robustus, H. modestus, H. lucassi and H. muelleri. 
H. kannemeyeri diners from all these in the structure of the petasma and 
thelycum. The latter most closely resembles that of H. fattahi Ramadan, 1938 
(as figured by Crosnier & Forest 1973, fig. 85g-h) but the petasma of H. fattahi 
differs markedly from the present species. 








Fig. 7. Hymenopenaeus kannemeyeri sp. nov. A. Carapace in lateral view. B. Anterior and 
posterior view of petasma. C. Thelycum. D. Telson. E. Eye. F. Pleopod 2, <$, appendix 

interna and appendix masculina. 



THE SOUTH AFRICAN MUSEUM'S MEIRING NAUDE CRUISES 29 

The species is named for S. X. Kannemeyer, of the Department of Marine 
Biology, South African Museum, in appreciation for his invaluable help in the 
Meiring Naude cruises. 

Genus Funchalia Johnson 

As Funchalia villosa and F. woodwardi have been confused in the past, the 
petasma and thelycum of each is figured, and all the South African Museum's 
holdings of both species are listed. 

Funchalia {Funchalia) villosa (Bouvier) 
Fig. 8C-D 
Funchalia {Funchalia) villosa: Burkenroad, 1936: 129. Crosnier & Forest, 1973: 296. 

Previous records 

Eastern and western North Atlantic; Caribbean; southern central South 
Atlantic. 

Material and South African records 

SAM-A15368 SM 18 1 $ CL 22,8 mm 

SAM-A15369 SM 33 1 $ CL 21,0 mm 

SAM-A 15370 SM 88 1 $ 1 $ CL 13,0 mm, 14,2 mm 

SAM-A15371 SM 111 1 S CL 18,4 mm 

SAM-A13195 off Natal, 30°30'S 31°45'E200 m 1 $ 

SAM-A13197 off Natal, 26°30'S 42°40'E 500 m 1 <? 

SAM-A13198 off Agulhas Bank, 37°10'S 21°50'E 500 m 1 <J 

SAM-A13236 off Natal, 31°44'S 44°35'E 500 m3^ 

Funchalia {Funchalia) woodwardi Johnson 
Fig. 8A-B 

Funchalia {Funchalia) woodwardi: Barnard, 1950: 609, fig. 112. Gordon & Ingle, 1956: 478 
Paulinose, 1974: 433. 

Previous records 

Eastern North Atlantic; Mediterranean; south-eastern South Atlantic. 

South African records 

SAM-A8481 off Table Bay, from stomach of stock fish 9 So 

SAM-A10914 off Cape Peninsula, from tunny stomach several $S & $$ 

SAM-A 10995 off Cape Peninsula, from tunny stomach 1 g 

SAM-A11025 off Dassen Island, from tunny stomach 25 $$ 19 ?$ 

SAM-A 1 1027 off Cape Peninsula, several SS & $$ 

SAM-A12014 off Dassen Island, 500 m2^2?? 

SAM-A 1 2482 Vema Seamount, from tunny stomach 60 immature S S & 9? 

SAM-A 13 196 off Cape Peninsula, 250 m 3 $$ 

SAM-A13237 off Cape Peninsula, 350 m7^7?? 

SAM-A13238 off Cape Peninsula, 500 m2^3?? 



30 



ANNALS OF THE SOUTH AFRICAN MUSEUM 







Fig. 8. 



B 

A. Funchalia woodwardi petasma. B. F. woodwardi thelycum. C. Funchalia villosa 
petasma. D. F. villosa thelycum. 



THE SOUTH AFRICAN MUSEUM'S ME1RING NAUDE CRUISES 31 

Family Sergestidae 

Serges tes {Sergia) talisman i Barnard 

Sergestes talismani Barnard, 1947: 384. 

Sergestes (Sergia) talismani: Crosnier & Forest, 1973: 325, figs llla-c, 1 12a-b. 

Previous records 

Cape Verde Islands; Portuguese Guinea; Gabon; Congo. 

Material 

See species list. 

Remarks 

This would seem to be the first record of the species from the Indian Ocean. 



CARIDEA 
Family Oplophoridae 

Acanthephyra armata Edwards 
Fig. 9B 

Acanthephyra armata: Bate, 1888: 744, pi. 125 (fig. 2). De Man, 1920: 61, pi. 6 (fig. 13). 
Chace, 1936: 27. 

Previous records 

West Indies; Bali Sea; Straits of Macassar; off Kei Islands; off Banda 
Island. 

Material 

SAM-A15154 SM38 4 $$ 4 $$ 

SAM-A15150 SM 58 3 <J(J 3 ?? 

SS CL 16,9 mm-29,0 mm RL 27,0 mm-37,5 mm TL 92,0 mm-137 mm 

$$ 24,5 mm-33,0 mm 34,7 mm-35,2 mm 121 mm-146 mm 

Acanthephyra indica Balss 
Fig. 9A 

Acanthephyra indica Balss, 1925: 264, fig. 34. Chace, 1936: 26. 
Acanthephyra sp. De Man, 1920: 68, pi. 6 (fig. 16). 

Previous records 

Between Sumatra and Ceylon; Banda Sea; off Ambon. 

Material 

See species list. 

3 $$ 1 ? 3 ovig. ?$ S CL 10,0 mm-10,2 mm $ CL 10,9 mm-13,1 mm 



32 



ANNALS OF THE SOUTH AFRICAN MUSEUM 




Fig. 9. A. Acanthephyra indica. B. Acanthephyra armata. 

Family Pasiphaeidae 

Eupasiphae gilesii (Wood-Mason & Alcock) 

Fig. 10B 

Parapasiphae {Eupasiphae) gilesii Wood-Mason & Alcock, 1893 : 166. Alcock, 1892: pi. 3 (fig. 8); 

1901 : 66. 
Eupasiphae gilesii: Holthuis, 1955: 36. Tirmizi, 1969: 214, figs 1-1. Foxton, 1970: 958. 
Eupasiphae gile si: Fisher & Goldie, 1961: 78. Crosnier & Forest, 1973: 150, fig. 44. 

Previous records 

Bermuda; Cape Verde Islands; Canary Islands; Madeira; Arabian Sea; 
Gulf of Oman ; Andaman Sea. 

Material 

SAM-A15151 SM 105 1£ CL + RL 32,4 mm TL 97 mm 



Pasiphaea sivado (Risso) 

Fig. 10A 

Pasiphae sivado: Wood-Mason & Alcock, 1893: 161. Alcock, 1892: pi. 3 (fig. 6); 1901: 59. 
Sivertsen & Holthuis, 1956: 29. Crosnier & Forest, 1973: 133. 



THE SOUTH AFRICAN MUSEUM'S MEWING NAUDE CRUISES 



33 





Fig. 10. A. Pasiphae sivado. B. Eupasiphae gilesi. 



34 



ANNALS OF THE SOUTH AFRICAN MUSEUM 



Previous records 

Eastern North Atlantic; Mediterranean; Red Sea; Bay of Bengal; Andaman 
Sea. 

Material 

SAM-15153 SM63 2 9? 
SAM-A15152 SM 85 1? 
9? CL 10,0 mm-14,5 mm 

Pasiphaea meiringnaudei sp. no v. 

Figs 11-14 
Description 

Carapace (Fig. 11) mid-dorsally carinate for almost its entire length. 
Rostrum a postfrontal triangular spine directed obliquely upward, only rarely 
reaching beyond anterior carapace margin. Anterior margin of rostrum straight, 
almost vertical, apparently more variable in females than in males (Fig. 13). 
Orbital angle a rounded lobe; branchiostegal spine directed outwards; supra- 
branchial ridge rounded, not reaching posterior carapace margin. 

Second to sixth abdominal segments dorsally keeled, not ending in a 
posterior spine; sixth segment with curved lateral ridge. Telson shorter than 




Fig. 1 1. Pasiphae meiringnaudei sp. nov. Holotype in lateral view, 



THE SOUTH AFRICAN MUSEUM'S MEIRING NAUDE CRUISES 



35 



uropodal endopod, dorsally grooved, distally forked, with seven spines on inner 
margin of sinus (Fig. 121). 

Antennular peduncle extending for two-thirds of scaphocerite (Fig. 11); 
stylocerite (Fig. 12G) broadening distally, twisted, apically acute, dorsolateral 
flagellum with about eighteen enlarged basal segments in male and female. 

Antennal base bearing ventrodistal spine. 




Fig. 12. Pasiphae meiringnaudei sp. nov. A, Anterior carapace and rostrum. B. Mandible. 

C. First maxilla. D. First maxilliped. E. Second maxilliped. F. Second maxilla. G. Antennular 

base. H. Antennal base. I. Telsonic apex. J. Pleopod 2, <$, appendix interna and appendix 

masculina. K. Pleopod 1, L L. Pleopod 1, cf, endopod. 



36 ANNALS OF THE SOUTH AFRICAN MUSEUM 

Third maxilliped just reaching to end of scaphocerite. 

First pereiopod overreaching scaphocerite by half length of palm plus 
fingers (Fig. 1 1); palm longer than fingers; merus with one to four ventral spines; 
basis with ventral distal triangular spine. 

Second pereiopod overreaching scaphocerite by half length of palm plus 
fingers ; palm equal in length to fingers ; carpus with broad ventral tooth ; merus 
with fifteen to twenty-one ventral spines; ischium with one spine, basis with four 
to seven spines (Fig. 14B). 

Third pereiopod overreaching anterior carapace margin by about half 
length of propodus plus dactylus; very slender; dactylus tapering; propodus one- 
third length of dactylus ; carpus very elongate, almost four times length of merus. 

Fourth pereiopod reaching to midpoint of carpus of third pereiopod. 

Fifth pereiopod longer than fourth pereiopod, reaching to end of carpus of 
third pereiopod. 

Endopod of first pleopod o* short, with triangular process on inner margin 
bearing very small denticles (Fig. 12L). 

Second pleopod <$ appendix masculina twice length of appendix interna, 
latter curved, with seven marginal setae (Fig. 12J). 

Uropodal endopod two-thirds length of exopod, latter with small terminal 
spine on outer margin. 

Material 

Holotype SAM-A15316 SM 107 ovig. ? CL 57,0 mm TL 166 mm 

SM 38 1 6* CL 39,9 mm TL 126 mm 

SM 58 4 $$ CL 35,5 mm-36,3 mm TL 109 mm-110 mm 

SM 107 5 ovig. $? 50,0 mm-58,0 mm 153 mm-168 mm 

SM 91 1 ? 42,0 mm 125 mm 

SM 28 1 $ 34,5 mm 108 mm 

SM40 2$$ 31,8 mm-42,4 mm 95-130 mm 









Fig. 13. Pasiphae meiringnaudei sp. nov. rostral variation. 

Top row, ovigerous ?$. Bottom row, $$. Bottom right, 

rostrum modified by ellobiopsid parasites. 



THE SOUTH AFRICAN MUSEUM'S M EI RING NAUDE CRUISES 



37 



Remarks 

J. C. Yaldwyn (National Museum of New Zealand) has kindly supplied 
most of the following information on the closely related species of Pasiphaea, 
from an as yet unpublished review of the genus by him and F. A. Chace Jr 
(Smithsonian Institution). The combination of three characters, viz. the distally 
forked telson, carinate carapace and abdomen, and basis of the second pereiopod 
ventrally spined, places the present species in a group with four previously 
described species. These are P. multidentata Esmark, P. tarda Kroyer, P. rath- 
bunae Stebbing, and P. barnardi Yaldwyn. The rostral shape of P. meiringnaudei 
differs from these four, but comes closest to P. barnardi. The finger-to-palm 
ratio of the first and second pereiopods, however, diners from Yaldwyn's species 
where the fingers of the first pereiopod are subequal to the palm and the fingers 
of the second pereiopod are distinctly longer than the palm. P. meiringnaudei 
overlaps with P. tarda (as P. princeps in Kemp 1910, pi. 4 (figs 1-7)) in the spine 
counts of the first and second pereiopods, but differs markedly in the rostral 
shape and in the relative lengths of the thoracic appendages. 

The species is named for the R/V Meiring Naude. 





Fig. 14. Pasiphae meiringnaudei sp. nov. A. First pereiopod, with chela further enlarged. 
B. Second pereiopod, with chela further enlarged. 



38 ANNALS OF THE SOUTH AFRICAN MUSEUM 

Family Pandalidae 

Heterocarpus dorsalis Bate 

Fig. 15A 

Heterocarpus dorsalis: De Man, 1920: 156, 171, pi. 15 (fig. 43). Barnard, 1950: 684, fig. 127a 

Previous records 

Off Durban; off East Africa; East Indies; Japan. 

Material 

See species list. 

7 S6 CL 20 mm-41 mm RL 35 mm-48 mm TL 98 mm-179 mm 

2$$ 27 mm-42 mm 35 mm-44 mm 126 mm-171 mm 

Remarks 

The ovigerous female of total length 171 mm is at least 20 mm longer than 
the largest ovigerous female recorded by De Man (1920). 

Heterocarpus laevigatus Bate 
Fig. 15B 

Heterocarpus laevigatus Bate, 1888: 636, pi. 112 (fig. 3). Stebbing, 1914: 40. De Man, 1920: 
154, 159, pi. 13 (fig. 37). Barnard, 1950: 684, fig. 127b. Crosnier & Forest, 1973: 195, fig. 
61c. 

Previous records 

Cape Verde Islands; Madeira; off East London; Arabian Sea; East Indies; 

Hawaii. 

Material 

See species list. 

2S3 CL 65 mm-74 mm RL 28 mm-38 mm TL 121 mm-160 mm 

3$$ 64 mm-105 mm 36 mm-51 mm 121 mm-200 mm 

Remarks 

The East London record was of an immature specimen measuring approxi- 
mately 63 mm TL. 

Heterocarpus tricarinatus Alcock & Anderson 
Fig. 15C 

Heterocarpus tricarinatus: De Man, 1920: 155, 161, pi. 13 (fig. 38), pi. 14 (fig. 38). Caiman, 
1939: 204. Barnard, 1950: 682, fig. 127c-d. 

Previous records 

Off East London; Arabian Sea; East Indies. 

Material 

SAM-A15264 SM 22 1 $ CL 35 mm RL 30 mm TL 135 mm 

ovig. $ 37 mm 32 mm 146 mm 



THE SOUTH AFRICAN MUSEUM'S MEIRING NAUDE CRUISES 



39 




'0™ 10 2 to 3'0 {.'Q 5 




|ltlt|llft{l!fl|llll| 



3'0 V0 5 j 



B 




|!il!;llll|i'l!i|iiri|iili|!lli|iiii|llllj!lll|ll!lj 
bmi. 1'0 2 ! 3'0 4 ! 5 ; 



Fig. 15. A. Heterocarpus dorsalis. B. Heterocarpus laevigatus. C. Heterocarpus tricarinatus. 



40 ANNALS OF THE SOUTH AFRICAN MUSEUM 

Remarks 

Using both Barnard's (1950) and De Man's (1920) keys to the genus 
Heterocarpus, H. tricarinatus is arrived at. The genus is obviously variable with 
regard to the rostral dentition, proportions of the pereiopodal dactyli, and the 
lengths of the carapace keels. This led Caiman (1939) to doubt the validity of 
this species, suggesting that it might be synonymous with H. gibbosus Bate. The 
variability of this latter species is well illustrated by De Man (1920). The present 
material could well be placed in Bate's species, but until this species complex 
is resolved, the lower carapace keel reaching back about two-thirds of the cara- 
pace length necessitates placing these two specimens in H. tricarinatus. 

Family Glyphocrangonidae 

Glyphocrangon dentatus Barnard 

Glyphocrangon gilesii var. dentata Barnard, 1926: 128. 
Glyphocrangon mabihissae Caiman, 1939: 217, fig. 8. 
Glyphocrangon dentarus Barnard, 1947 (typ. err.): 387. 
Glyphocrangon dentatus Barnard, 1950: 722, fig. 134h. 

Previous records 

Off Mozambique, 540 m; off Zanzibar, 640-658 m. 

Material 

8 6*6* 1 ? 8 ovig. $$, see species list. 

6*c? CL 16,5 mm-19,8 mm RL 17,8 mm-22 mm TL 73 mm-85 mm 
ovig. $$ 19,1 mm-23 mm 20,8 mm-27,5 mm 83 mm-102 mm 

Remarks 

Only slight differences are apparent between Caiman's specimens from 
Zanzibar and the present material. These differences are to some extent probably 
due to individual variation, as well as to variations between populations. The 
spine on the midpoint of the posterior antennal carina is present in some of the 
present specimens, absent in others, while the row of small tubercles just lateral 
to the submedian carina seen in the Zanzibar specimens is represented by one 
or two almost indistinct tubercles. Although the rostral length always exceeds 
the carapace length, the ratio is variable. 

Glyphocrangon regalis Bate 
Fig. 16 
Glyphocrangon regalis Bate, 1888: 517, pi. 93 (figs 3-4). De Man, 1920: 220. 

Description 

Rostrum curved ventrally, with apex upturned, slightly less than two-thirds 
carapace length, with pair of lateral rostral teeth above eyes, second pair of 
spines in postorbital position. Antennal spine strong, directed forward, separated 



THE SOUTH AFRICAN MUSEUM'S MEIRING NAUDE CRUISES 



41 



by short, curved carapace margin from longer branchiostegal spine directed 
anteroventrally. Anterior submedian carinae consisting of seven elongate 
tubercles. Posterior submedian carinae consisting of four elongate tubercles. 
Row of small, rounded tubercles median to both anterior and posterior sub- 
median carinae. Small median spine at base of rostrum flanked by two postero- 
lateral spines. Anterior antennal carina forming wing-like flattened structure 
ending in outward-pointed strong spine. Posterior antennal carina situated some 
distance above anterior antennal carina, ending anteriorly in outwardly directed 
spine, not as strong as that of anterior carina ; at about midpoint, a slight indica- 
tion of a notch, never strong enough to be called a spine or tubercle. Anterior 
intermediate carina not well defined, consisting of three or four large irregularly 
placed tubercles. Posterior intermediate carina consisting of four or five elongate 
tubercles. Area between anterior submedian and anterior intermediate carinae 
with two rows of small tubercles; three rows of tubercles between posterior 
submedian and posterior intermediate carinae. Area between anterior antennal 
carina and lateral groove carrying a scattering of small rounded tubercles. Areas 
between posterior antennal and posterior lateral carinae also carrying irregular 




Fig. 16. Glyphocrangon regalis. 



42 ANNALS OF THE SOUTH AFRICAN MUSEUM 

scattering of rounded tubercles. Anterior and posterior lateral carina entire, not 
tuberculate. Sublateral carina only distinct in posterior region, consisting of a 
foveolate ridge. Region between sublateral carina and margin bearing a foveolate 
band. 

Abdomen (excluding telson) equal in length of carapace plus rostrum. 
Median carina of abdominal segments strong, only on first segment forming a 
forwardly directed spine. Pleura of first segment considerably shorter than second, 
with forwardly directed spine. Pleura of second segment ventrally bispinose, 
spines of equal strength. Pleura of third segment ventrally bispinose, anterior 
spine longer than posterior. Pleura of fourth segment ventrally bispinose, spines 
subequal. Pleura of fifth segment bispinose, spines directed posteroventrally. 
Sixth abdominal segment ending posteroventrally in strong spine. Telson with 
median spine at base, elongate triangular, with strong dorsolateral keel, apex 
acute, flexed slightly dorsally. 

Eyes well developed, with large black corneas, small spinose tubercle on 
inner distal margin of eye-stalk. 

Scaphocerite ovate, fringed with setae, lacking any spine on inner margin. 

Second pereiopods unequal, left shorter and slightly squatter than right, 
carpus of right (30-31 segments) considerably longer than left (19 segments), 
chela markedly smaller. 

Third pereiopod dactylus lanceolate, oval in cross section, one-third length 
of propodus. 

Fourth pereiopod dactylus slightly more than one-third length of propodus, 
dorsally grooved. 

Fifth pereiopod dactylus similar to fourth. 

Previous records 

Banda Island; off Makassar; Kermadec Island; Fiji; Paternoster Island. 

Material 

5 SS 7 9? 10 ovig. 99, see species list. 

<$<$ CL 20,5 mm-29,9 mm RL 16,5 mm-19,9 mm TL 82,5 mm-108 mm 
ovig. ?? 29,3mm-34,9mm 17,8 mm-21,0mm 108mm-128mm 

Remarks 

The species recorded by Wood-Mason (1891) as G. investigatoris and 
G. investigatoris var. andamanensis, was stated by De Man (1920) to be possibly 
synonymous with G. regalis. Wood-Mason's species, however, shows some 
differences, e.g. in the degree of 'flare' of the spine of the postantennal carina, the 
possession of midline tubercles on the rostrum which together suggest that this 
is not the same species. Examination of the 'Challenger' types of G. regalis reveal 
almost no differences from the present material. 



THE SOUTH AFRICAN MUSEUM'S ME/RING NAUDE CRUISES 43 

ACKNOWLEDGEMENTS 

In addition to the people and institutions thanked in the introductory paper 
of this series, I should like to thank Mr S. X. Kannemeyer of the South African 
Museum for the photographs, Dr R. B. Manning of the Smithsonian Institution 
for the loan of Pasiphaea material, Dr J. Yaldwyn of the National Museum of 
New Zealand and Dr F. A. Chace Jr of the Smithsonian Institution for informa- 
tion on Pasiphaea, and Prof. J. H. Day of the University of Cape Town and 
Dr J. Yaldwyn for reading the manuscript and for their many useful criticisms. 

REFERENCES 

Alcock, A. 1892. Crustacea Part I. Illustrations of the Zoology of the Investigator Plates 1-5. 

Calcutta: Superintendent of Government Printing. 
Alcock, A. 1899. Crustacea Part VII. Illustrations of the Zoology of the Investigator Plates 

36-45. Calcutta: Superintendent of Government Printing. 
Alcock, A. 1901. A descriptive catalogue of the Indian deep-sea Crustacea Decapoda Macrura 

and Anomala, in the Indian Museum. Calcutta: Trustees of the Indian Museum. 
Alcock, A. & Anderson, A. R. S. 1899. Natural history notes from H.M. Royal Indian 

Marine Survey Ship 'Investigator', Commander T. H. Heming, R.N., commanding. 

Series III, no. 2. An account of the deep-sea Crustacea dredged during the surveying 

season 1897-98. Ann. Mag. nat. Hist. (7) 3: 278-282. 
Alcock, A. & McArdle, A. F. 1901. Crustacea Part IX. Illustrations of the Zoology of the 

Investigator Plates 49-55. Calcutta: Superintendent of Government Printing. 
Balss, H. 1925. Macrura der Deutschen Tiefsee-Expedition. 2. Natantia, Teil A. Wiss. Ergebn. 

dt. Tiefsee-Exped. 'Valdivia" 20: 221-315. 
Barnard, K. H. 1926. Report on a collection of Crustacea from Portuguese East Africa. 

Trans. R. Soc. S. Afr. 13: 119-130. 
Barnard, K. H. 1947. Descriptions of new species of South African decapod Crustacea, with 

notes on synonymy and new records. Ann. Mag. nat. Hist. (11) 13: 361-392. 
Barnard, K. H. 1950. Descriptive catalogue of South African decapod Crustacea (crabs and 

shrimps). Ann. S. Afr. Mus. 38: 1-837. 
Berry, P. F. & Plante, R. 1973. Revision of the spiny lobster genus Palinurus in the south- 
west Indian Ocean. Trans. R. Soc. S. Afr. 40: 373-380. 
Bate, C. S. 1888. Report on the Crustacea Macrura dredged by HMS Challenger during the 

years 1873-1876. Rep. Voy. Challenger, 1873-1876 24: 1-942. 
Burkenroad, M. D. 1936. The Aristaeinae, Solenocerinae and pelagic Penaeinae of the 

Bingham Oceanographic Collection. Bull. Bingham oceanogr. Coll. 5 (2): 1-151. 
Burucovsky, R. N. & Romensky, L. L. 1972. On rostrum changeability of Aristeus varidens 

Holthuis (Decapoda, Penaeidae). At/. Nauchno-Issled. Inst. Rybn. Khoz. Okeanogr. 42: 

156-161. 
Calman, W. T. 1939. Crustacea: Caridea. Scient. Reps. John Murray Exped. 6: 183-223. 
Chace, F. A. 1936. Revision of the bathypelagic prawns of the family Acanthephyridae, with 

notes on a new family, Gomphonotidae. J. Wash. Acad. Sci. 26: 24-31. 
Crosnier, A. & Forest, J. 1973. Faune Tropicale. XIX. Les Crevettes profondes de FAtlantique 

Oriental Tropical. Paris: O.R.S.T.O.M. 
Fisher, L. R. & Goldie, E. H. 1961. New records of two deep-sea decapods. Crustaceana 2: 

78-79. 
Foxton, P. 1970. The vertical distribution of pelagic decapods (Crustacea: Natantia) collected 

on the SOND cruise 1965. I. The Caridea. J. mar. biol. Assoc. U.K. 50: 939-960. 
George, R. W. & Grindley, J. R. 1964. Projasus—a new generic name for Parker's crayfish, 

Jasus parkeri Stebbing (Palinuridae: 'Silentes'). J. Proc. R. Soc. West. Aust. 47: 87-90. 
Gordon, I. & Ingle, R. W. 1956. On a pelagic penaeid prawn, Funchalia woodwardi Johnson, 

new to the British fauna. /. mar. biol. Assoc. U.K. 35: 475-481. 
Hall, D. N. F. 1962. Observations on the taxonomy and biology of some Indo-West Pacific 

Penaeidae. Col. Off. Fish. Pubis 17: 1-230. 



44 ANNALS OF THE SOUTH AFRICAN MUSEUM 

Holthuis, L. B. 1955. The recent genera of the Caridean and Stenopodidean shrimps (Class 

Crustacea, Order Decapoda, Supersection Natantia) with keys for their determination. 

Zool. Verhandl. 26: 1-157. 
Kemp, S. 1910. The Decapoda Natantia of the coasts of Ireland. Scient. Invest. Fish. Brch Ire. 

1908(1): 1-190. 
Kensley, B. F. 1969. Decapod Crustacea from the south-west Indian Ocean. Ann. S. Afr. Mus. 

52: 149-181. 
Kensley, B. F. 1971a. The Family Sergestidae in the waters around southern Africa (Crustacea, 

Decapoda, Natantia). Ann. S. Afr. Mus. 57: 215-264. 
Kensley, B. F. 19716. The genus Gennadas in the waters around southern Africa. Ann. S. Afr. 

Mus. 57: 271-294. 
Louw, E. M. 1977. The South African Museum's Meiring Naude cruises. Part 1. Station data 

1975, 1976. Ann. S. Afr. Mus. 72: 147-159. 
Man, J. G. de. 1911. The Decapoda of the Siboga Expedition. Part I. Family Penaeidae. 

Siboga Exped. monogr. 39A: 1-131. 
Man, J. G. de. 1920. The Decapoda of the Siboga Expedition. Part IV. Families Pasiphaeidae, 

Stylodactylidae, Hoplophoridae, Nematocarcinidae, Thalassocaridae, Pandalidae, Psalido- 

podidae, Gnathophyllidae, Processidae, Glyphocrangonidae, and Crangonidae. Siboga 

Exped. monogr. 39 A3: 1-318. 
Paulinose, V. T. 1974. Decapod Crustacea from the International Indian Ocean Expedition: 

The species of Funchalia (Penaeidae) and their post-larvae. /. nat. Hist. 8: 433-443. 
Ramadan, M. M. 1938. Crustacea, Penaeidae. Scient. Reps John Murray Exped. 5: 35-76. 
Roberts, T. W. & Pequegnat, W. E. 1970. Deep-water decapod shrimps of the Family 

Penaeidae. In: Pequegnat, W. E. & Chace, F. A. Texas A &M University Oceanographical 

Studies 1 . Contributions on the Biology of the Gulf of Mexico. Houston, Texas : Gulf 

Publishing Co. 
Sivertsen, E. & Holthuis, L. B. 1956. Crustacea Decapoda (the Penaeidae and Stenopodidae 

excepted). Rep. scient. Results Michael Sars N. Atlant. Deep Sea Exped. 5 (12): 1-54. 
Stebbing, T. R. R. 1914. South African Crustacea. Ann. S. Afr. Mus. 15: 1-55. 
Tirmizi, N. 1969. Eupasiphae gilesii (Wood-Mason, 1892) from the northern Arabian Sea 

(Decapoda, Caridea). Crustaceana 16: 213-218. 
Wood-Mason, J. & Alcock, A. 1891. On the results of deep-sea dredging during the season 

1890-91. Ann. Mag. nat. Hist. (6) 8: 268-286. 
Wood-Mason, J. & Alcock, A. 1893. On the results of deep-sea dredging during the season 

1890-91. Ann. Mag. nat. Hist. (6) 11: 161-171. 



6. SYSTEMATIC papers must conform to the International code of zoological nomenclature 
(particularly Articles 22 and 51). 

Names of new taxa, combinations, synonyms, etc., when used for the first time, must be 
followed by the appropriate Latin (not English) abbreviation, e.g. gen. nov., sp. nov., comb, 
nov., syn. nov., etc. 

An author's name when cited must follow the name of the taxon without intervening 
punctuation and not be abbreviated; if the year is added, a comma must separate author's 
name and year. The author's name (and date, if cited) must be placed in parentheses if a 
species or subspecies is transferred from its original genus. The name of a subsequent user of 
a scientific name must be separated from the scientific name by a colon. 

Synonymy arrangement should be according to chronology of names, i.e. all published 
scientific names by which the species previously has been designated are listed in chronological 
order, with all references to that name following in chronological order, e.g.: 

Family Nuculanidae 

Nuculana (Lembulus) bicuspidata (Gould, 1 845) 

Figs 14-15A 
Nucula (Leda) bicuspidata Gould, 1845: 37. 
Leda plicifera A. Adams, 1856: 50. 

Laeda bicuspidata Hanley, 1859: 118, pi. 228 (fig. 73). Sowerby, 1871 : pi. 2 (fig. 8a-b). 
Nucula largillicrti Philippi, 1861 : 87. 
Leda bicuspidata: Nickles, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9. 

Note punctuation in the above example: 

comma separates author's name and year 

semicolon separates more than one reference by the same author 

full stop separates references by different authors 

figures of plates are enclosed in parentheses to distinguish them from text-figures 

dash, not comma, separates consecutive numbers 

Synonymy arrangement according to chronology of bibliographic references, whereby 
the year is placed in front of each entry, and the synonym repeated in full for each entry, is 
not acceptable. 

In describing new species, one specimen must be designated as the holotype; other speci- 
mens mentioned in the original description are to be designated paratypes; additional material 
not regarded as paratypes should be listed separately. The complete data (registration number, 
depository, description of specimen, locality, collector, date) of the holotype and paratypes 
must be recorded, e.g. : 

Holotype 

SAM-A 13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King's Beach 
Port Elizabeth (33°51'S 25°39'E), collected by A. Smith, 15 January 1973. 

Note standard form of writing South African Museum registration numbers and date. 

7. SPECIAL HOUSE RULES 

Capital initial letters 

(a) The Figures, Maps and Tables of the paper when referred to in the text 

e.g. '. . . the Figure depicting C. namacolus . . .'; '. . . in C. namacolus (Fig. 10) . . .' 

(b) The prefixes of prefixed surnames in all languages, when used in the text, if not preceded 
by initials or full names 

e.g. Du Toit but A. L. duToit; Von Huene but F. von Huene 

(c) Scientific names, but not their vernacular derivatives 
e.g. Therocephalia, but therocephalian 

Punctuation should be loose, omitting all not strictly necessary 
Reference to the author should be expressed in the third person 
Roman numerals should be converted to arabic, except when forming part of the title of a 

book or article, such as 

'Revision of the Crustacea. Part VIII. The Amphipoda.' 
Specific name must not stand alone, but be preceded by the generic name or its abbreviation 

to initial capital letter, provided the same generic name is used consecutively. 
Name of new genus or species is not to be included in the title: it should be included in the 

abstract, counter to Recommendation 23 of the Code, to meet the requirements of 

Biological Abstracts. 



BRIAN KENSLEY 

THE SOUTH AFRICAN MUSEUM'S 
MEIRING NAUDE CRUISES 

PART 5 
CRUSTACEA, DECAPODA, REPTANTIA 

AND NATANTIA 



50U 7^^ 

VOLUME 74 PART 3 NOVEMBER 1977 



ISSN 0303-2515 



MUS. COMP. ZOOU 
LIBRARY 

'JAN 3 1 1978 

HARVARD 
UNIVERSITY 



ANNALS 




INSTRUCTIONS TO AUTHORS 

1. MATERIAL should be original and not published elsewhere, in whole or in part. 

2. LAYOUT should be as follows: 

(a) Centred masthead to consist of 

Title : informative but concise, without abbreviations and not including the names of new genera or species 

Author's(s') name(s) 

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(b) Abstract of not more than 200 words, intelligible to the reader without reference to the text 

(c) Table of contents giving hierarchy of headings and subheadings 

(d) Introduction 

(e) Subject-matter of the paper, divided into sections to correspond with those given in table of contents 

(f ) Summary, if paper is lengthy 

(g) Acknowledgements 
(h) References 

(i) Abbreviations, where these are numerous 

3. MANUSCRIPT, to be submitted in triplicate, should be typewritten and neat, double spaced 
with 2,5 cm margins all round. First lines of paragraphs should be indented. Tables and a list of 
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Major headings of the paper are centred capitals; first subheadings are shouldered small 
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figures is not set in type and should be in lower-case letters. 

The number of the figure should be lightly marked in pencil on the back of each illustration. 

5. REFERENCES cited in text and synonymies should all be included in the list at the end of 
the paper, using the Harvard System {ibid., idem, be. cit., op. cit. are not acceptable): 

(a) Author's name and year of publication given in text, e.g. : 

'Smith (1969) describes . . .' 

•Smith (1969: 36, fig. 16) describes . . .' 

*As described (Smith 1969a, 19696; Jones 1971)' 

'As described (Haughton & Broom 1927) . . .' 

'As described (Haughton et al. 1927) . . .' 

Note: no comma separating name and year 

pagination indicated by colon, not p. 

names of joint authors connected by ampersand 

et al. in text for more than two joint authors, but names of all authors given in list of references. 

(b) Full references at the end of the paper, arranged alphabetically by names, chronologically 
within each name, with suffixes a, b, etc. to the year for more than one paper by the same 
author in that year, e.g. Smith (1969a, 19696) and not Smith (1969, 1969a). 

For books give title in italics, edition, volume number, place of publication, publisher. 

For journal article give title of article, title of journal in italics (abbreviated according to the World list o 
scientific periodicals. 4th ed. London: Butter worths, 1963), series in parentheses, volume number, part 
number (only if independently paged) in parentheses, pagination (first and last pages of article). 

Examples (note capitalization and punctuation) 

Bullough, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan. 

Fischer, P.-H. 1948. Donnees sur la resistance et de le vitalite des mollusques. /. Conch., Paris 88: 100-140. 

Fischer, P.-H., Duval, M. & Raffy, A. 1933. Etudes sur les echanges respiratoires des littorines. Archs 

Zool. exp. gen. 14: 627-634. 
Kohn, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon. 

Ann. Mag. nat. Hist. (13) 2: 309-320. 
Kohn, A. J. 19606. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean. 

Bull. Bingham oceanogr. Coll. 17 (4): 1-51. 
Thiele, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. In: Schultze, L. Zoologische 

und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Sud-Afrika 4 : 269-270. 

Jena: Fischer. Denkschr. med.-naturw. Ges. Jena 16: 269-270. 

{continued inside back cover) 



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Court Road, Wynberg, Cape 



In Suid-Afrika gedruk deur 
Die Rustica-pers, Edms., Bpk., 
Courtweg, Wynberg, Kaap 



ILLUSTRATED GENERIC KEY TO 
SOUTH AFRICAN CONTINENTAL OSTRACODA 

By 

K. G. McKenzie 

Riverina College of Advanced Education, 
Wagga Wagga, Australia 

(With 24 figures) 
[MS. accepted 7 June 1977] 

ABSTRACT 

An illustrated generic key to South African continental ostracodes is provided together 
with the diagnoses of two new cypridopsinid genera, Sarscypridopsis and Kapcypridopsis. 





PAGE 


Introduction . 


. 45 


Systematics 


. 46 


Key 


. 49 


Acknowledgements . 


. 53 


References 


. 54 


Appendix 


. 55 



INTRODUCTION 

In a previous paper (McKenzie \91\a) the known continental and mainly 
freshwater Ostracoda of South Africa were listed together with a complete 
bibliography of such ostracode studies in the Republic. 

This paper complements that previous one by providing an illustrated key 
to the known ostracode genera of South Africa. The key is meant to be used with 
reference to illustrations in earlier papers by Sars (1896, \924a, 19246) and Rome 
(1965) as well as to the illustrations in this text. 

Ostracoda are microscopic crustaceans with a bivalve shell which usually 
completely covers the body when it is closed. This shell is commonly called the 
carapace. The ostracode body has 5-7 pairs of limbs— 7 pairs in freshwater 
taxa— and, usually, a furca at the posterior, the rami of which attach to paired 
chitin supports. From the anterior, the limbs are termed as follows: antennule; 
antenna; mandible; maxillule; and the first through to the third thoracic limbs 
(P I to P III). The furca normally carries two claws and two bristles on each 
ramus but can be strongly modified, e.g. into a whip-like seta in Cypridopsinae. 
Sometimes, as in Darwinula, it is absent. 

The furca in the genus Gesa is described in the key as having two claws 
only. This follows the original description (Hartmann 1957). However, this 

45 
Aim. S. Afr. Mus. 74 (3), 1977: 45-103, 24 figs. 



46 ANNALS OF THE SOUTH AFRICAN MUSEUM 

original description was of a juvenile form not of an adult. It is probable that 
the furca in an adult Gesa will be found to carry the normal complement of 
claws and bristles on each ramus. 

Some characters which are illustrated in the Figures are not normally 
recorded for ostracodes. These include the a, p and y bristles on the mandible 
palp which have been so named by Danielopol (personal communication 1970) 
and which appear to have some utility in taxonomy (e.g. compare Figures 6 
and 14). Similarly, rake-like organs are very rarely illustrated but may also 
prove to be useful taxonomic characters when known for more genera 
(McKenzie 19716). 

There is also a special terminology for characters of the ostracode carapace. 
These include: adductor muscle scar patterns (Fig. 1 et seq.); marginal or radial 
pore canals; normal pore canals which can be either simple and open or sieve- 
like (Fig. 1); and various types of hingement. The hingement terminology used 
here is that used in the American Treatise on Ostracoda (Moore ed. 1961) and 
is illustrated in Figure 1. 

Marginal pore canals are used to distinguish between genera in the tribe 
Stenocypridini. In particular, the genus Chrissia Hartmann, 1957 is distinguished 
by its very short marginal pore canals, whereas the species which were ascribed 
to Parastenocypris (McKenzie 1971a) have well-developed branching marginal 
pore canals (Figs 15-17). 

In this paper the new taxa comprise two new cypridopsinid genera which 
were foreshadowed earlier (McKenzie 1971 o), namely : the genus Sarscypridopsis\ 
and a new genus from Table Mountain, Kapcypridopsis. These new genera are 
described in the systematics section below. 

SYSTEMAT1CS 

Family Cyprididae 

Subfamily Cypridopsinae 

Tribe Cypridopsini 

Genus Kapcypridopsis gen. nov. 

Figs 6-7 
Type species 

Kapcypridopsis barnardi sp. nov. 

Derivation of name 

Kap = Cape (Afrikaans), barnardi for the late Dr K. H. Barnard, a 
tireless worker on South African crustaceans and molluscs and former director 
of the South African Museum. 

Diagnosis 

In lateral view, carapace small ; subreniform ; smooth ; greenish or yellowish ; 
valves subequal; dorsal margin convex; ventral margin inflexed medially; 



ILLUSTRATED GENERIC KEY TO SOUTH AFRICAN CONTINENTAL OSTRACODA 47 

anterior margin slightly more broadly rounded than the posterior; valve height 
slightly more than half valve length. In dorsal view, elliptical. Internally, inner 
lamellae broad anteriorly and posteriorly; inner margin regularly curved; 
vestibule large; line of concrescence following the outer margin; marginal pore 
canals short, fairly numerous; normal pore canals open and simple; muscle scar 
pattern including 4 adductor and 2 mandibular scars (sometimes individual 
adductor scars are divided and the adductor group appears to consist of 5 or 
6 scars); hingement of simple ridge and groove type. , 

Antennule 7-segmented; length ratios of the 5 proximal segments are 
20 : 13 : 10 : 10 : 15 (or 15 : 10 : 8 : 8 : 11). Antenna 4-segmented, length ratios 
of the 3 proximal segments are 7:4: 1 ; 'natatory' setae 6 in number, short, 
reaching about the middle of the second endopod segment. Mandible coxa 
normal; respiratory epipodial plate with 8 Strahlen; palp 4-segmented; a, j3 
and y bristles as illustrated. Maxillule respiratory epipod with 4 downwards- 
directed setae plus 17 Strahlen; lobes and palp elongate; palp 2-segmented, 
3rd lobe with two smooth Zahnborsten. PI epipod with 4 Strahlen; palp 
strongly dimorphic as is usual in cypridaceans with 3 terminal bristles in females 
but modified into a clasping organ in males. P II normal, with a powerful 
terminal claw. P III with a short terminal segment which bears a fairly short 
reflexed seta. Furca strongly modified, whip-like. Rake-like organs each with 
about 10 teeth. Eye with 2 ommatophores. In males, other dimorphic characters 
include a Zenkers organ with 13 whorls; and hemipenis as illustrated (Fig. 7: 7). 

Dimensions 

Length, about 0,6 mm; height, about 0,35 mm. 

Material 

Small population of six individuals including mature males and ovigerous 
females. Some forms were greenish, others yellowish, when collected. 
Syntypes: South African Museum catalogue number SAM-A 15451. 

Locality 

Top of Table Mountain, in small pool about 400 m from the cable car 
terminus; Cape Town, Cape Province, the Republic of South Africa. 

Collector and data collected 

K. G. McKenzie; September 1970. 

Discussion 

The regularly subreniform shape of this taxon is sufficient to distinguish it 
from Plesiocypridopsis Rome, Cypridopsis Brady, and Sarscypridopsis gen. nov. 
since Plesiocypridopsis and Sarscypridopsis are subtriangular in shape while 
Cypridopsis is subovate and this new genus is subreniform. Also, unlike these 
genera, Kapcypridopsis has broad anterior and posterior inner lamellae. 



48 ANNALS OF THE SOUTH AFRICAN MUSEUM 

Genus : Sarscypridopsis gen. nov. 

Fig. 8 
Type species 

Sarscypridopsis gregaria (Sars) 1896. 

Derivation of name 

For G. O. Sars, pioneer worker on South African Ostracoda. 

Diagnosis 

In lateral view, carapace small-medium; subtriangular; surface pitted and 
frequently hirsute; often greenish to brown; valves subequal; dorsal margin 
strongly convex; ventral margin inflexed medially; anterior margin rounded, 
tending to narrow anteroventrally; posterior margin more broadly rounded 
than the anterior margin; valve height usually well over half valve length. In 
dorsal view elongate subovate moderately broad (breadth about half the length); 
greatest breadth behind the middle. Internally, inner lamellae moderately broad 
anteriorly and narrow posteriorly; inner margin regularly curved, vestibule 
large, line of concrescence following the outer margin; marginal pore canals 
short, numerous ; normal pore canals simple, open ; central muscle scar pattern 
of 4 adductors and 2 mandibulars; hingement of simple ridge and groove type. 

Antennule 7-segmented; antenna 4-segmented; natatory setae extending 
well beyond the tips of the terminal claws; mandible normal, a, fi and y bristles 
as illustrated (Fig. 8: 2); maxillule palp 2-segmented, elongate-cylindrical; 
3rd lobe with 2 smooth Zahnborsten; PI epipod with 2 Strahlen; endopod 
modified in males into a clasping organ which is dissimilar between the right 
and left limbs (Fig. 8: 10, 13); P II normal with powerful terminal claw; 
PHI with a short reflexed seta; furca whip-like. In males, other dimorphic 
characters include a Zenkers Organ with 10-15 whorls; and hemipenis as 
illustrated. 

Discussion 

Of the three species in the original designation of Cypridopsis by Brady and 
Norman in 1889, one, Cypridopsis vidua (O. F. Muller, 1776), is the type species; 
C. villosa (Jurine, 1 806) was made the type species of Cypridopsella Kaufmann, 
1900, which is now synonymised with Potamocypris Brady, 1868, and C. aculeata 
Costa, 1847, was included by Sars (1924a: 160) with the South African species 
of Cypridopsis in his understanding of the genus at that time and indeed was 
made its type. However, C. vidua has priority as the type species of 
Cypridopsis s.s. which means that a different generic name is required for the 
African and possibly one European species (note that the type material of 
Costa's species has been lost and his original figures and descriptions were poor). 

The tribe Cypridopsini is differentiated from other cypridopsine ostracodes 
by the fact that it possesses an elongate maxillule palp and lobes, the palp being 



ILLUSTRATED GENERIC KEY TO SOUTH AFRICAN CONTINENTAL OSTRACODA 49 

further characterized by an elongate second segment. This feature is in contra- 
distinction to the broad palp, especially at the second segment, of Potamocypris 
and its related cypridopsine genera. Considered in this sense, Cypridopsini 
include the genera Cypridopsis, Plesiocypridopsis, Neocypridopsis, Kapcypridopsis, 
the present new genus Sarscypridopsis, and two Korean subspecies (McKenzie 
1972) which appear to be generically distinct from these others although similar 
in some features to Kapcypridopsis. 

This genus differs from Cypridopsis in that the shape is subtriangular and 
moderately broad rather than subovate and broad; the anterior inner lamellae 
are narrower, the ventral margin of the right valve is not denticulate; and there 
are only up to two Strahlen on the epipod of the P I. It differs from Plesio- 
cypridopsis in lacking the elongation of male P I palps and the characteristic 
upturned anterior hemipenes of that genus. Kapcypridopsis is subreniform in 
shape with broader anterior and posterior inner lamellae and short antennal 
'natatory' setae, differing in all these characters from Sarscypridopsis. The 
Korean taxa differ somewhat in shape especially posteriorly and have broad 
anterior and posterior inner lamellae and short antennal 'natatory' setae. The 
genus Neocypridopsis differs from Sarscypridopsis in general shape and also in 
the setation of the terminal segment of the P III, the shapes of the clasping 
palps of the male P I and in possessing an upturned not a downturned anterior 
hemipenis. 

Of the cypridopsinid species listed by Sars (1924c) aculeata Costa, 1847, 
spinifera Sars, 1924c, and echinata Sars, 1924c, form a subgroup characterized 
by an armature of spines as well as hairs on the shell surface. These species are at 
present classed with Sarscypridopsis but may merit transfer to a new category 
when aculeata Costa, 1847, is better understood. Apart from this latter species, 
which has been recorded from Iceland, Europe, central Asia, the Americas and 
Africa, Sarscypridopsis has a southern distribution. 

KEY 

1. All three pairs of thoracic legs more or less pediform and non- 
reflexed . . . . . . . . . . . . . . . . 2. 

First pair of thoracic legs non- or subpediform, strongly dimorphic; 
second pair pediform ; third pair strongly reflexed .. Cyprididae 7. 

2. Shell small; smooth; elongate-ovate in lateral view; adductor-muscle 
scar pattern feather-shaped ; first pair of thoracic legs (P I) sub- 
pediform ; males not reported since 1 870 

Darwinulidae, Darwinula (Fig. 1) 
Shell small to medium; surface usually ornamented; subrectangular, 
subreniform, subtrapezoidal or subrhomboidal in lateral view; 
adductor-muscle scar pattern of four discrete scars in a subvertical 
row; all three pairs of thoracic limbs pediform; typically bisexual 

Cytheridae 3. 



50 ANNALS OF THE SOUTH AFRICAN MUSEUM 

3. Shells small or medium; ornamented; subrectangular to subreniform; 
radial pore canals few, straight; normal pore canals simple; hinges 
modified lophodont . . . . . . . . Limnocytherinae 4. 

Shells small or medium; ornamented; subrectangular, subtrapezoidal 

or subrhomboidal; radial pore canals few or numerous, usually 
straight; normal pore canals of sieve type; hinge types merodont, 
entomodont or gonglyodont . . Cytherideinae, Loxoconchinae 5. 

4. Shell small; subrectangular to subreniform; non-dimorphic Limnocy there 
Shell medium; subrectangular; strongly dimorphic (females very 
broad, but males slender, posteriorly) . . . . Gomphocythere (Fig. 2) 

5. Shell small to medium; subrectangular or subtrapezoidal; radial pore 
canals few or numerous; hinge type merodont or entomodont 

Cytherideinae 6. 
Shell small; subrhomboidal; radial pore canals few, straight; hinge 
type gonglyodont . . . . . . Loxoconcha (Fig. 1) 

6. Shell small-medium; subtrapezoidal; radial pore canals few; hinge 
type merodont . . . . . . . . . . . . Perissocytheridea 

Shell medium; subrectangular; radial pore canals numerous; hinge 
type entomodont . . . . . . . . . . . . Cyprideis (Fig. 1) 

7. Shell medium; subrectangular; surface pitted, dorsomedial sulci 
present; palps of first pair of thoracic legs two-segmented 

Ilyocypridinae, Ilyocypris (Fig. 3) 
Shell differing from this; palps of first pair of thoracic legs non- 
segmented . . . . . . . . . . . . . . . . 8. 

8. Third thoracic limb bearing at least one long reflexed terminal seta; 
Zenkers Organ (males only) with 5-6 whorls 

Paracypridinae, Candoninae and Cyclocypridinae 9. 
Third thoracic limb with short reflexed terminal seta; Zenkers Organ 
(males only) with more than 6 whorls . . . . . . . . . . 12. 

9. Shell medium, subreniform or elongate subtriangular in lateral view; 
elliptical in dorsal view, antennal natatory setae poorly developed or 
absent . . . . . . . . Paracypridinae, Candoninae 10. 

Shell small-medium-, subquadrate in lateral view; antennal natatory 
setae well developed (extending to the terminal claws or beyond) 

Cyclocypridinae 1 1 . 

10. Shell medium, elongate-subtriangular in lateral view; furca with a 
posterior bristle, furcal claws without spurs Paracypridinae, Paracypris 
Shell medium, subreniform in lateral view; furca lacking a posterior 
bristle, furcal claws with spurs . . . . Candoninae, Candonopsis 

11. Shell small-medium; right valve denticulate ventrally especially on 
antero ventral margin; compressed in dorsal view; furca normal 

Cyclocypridinae, Physocypria (Figs 3-4) 
Shell small; non-denticulate ventrally; broadened in dorsal view, furca 
normal . . . . . . . . . . Cyclocypridinae, Cyclocypris 



ILLUSTRATED GENERIC KEY TO SOUTH AFRICAN CONTINENTAL OSTRACODA 51 

12. Shells small to very large; furca with at least two terminal claws . . 19. 
Shells small to medium; furca whip-like. . .. Cypridopsinae 13. 

13. Shell with distinctive pustulose surface ornamentation and charac- 
teristic lunette-bearing inner lamellae; maxillule 3rd lobe with at least 
four Zahnborsten . . . . . . . . . . Oncocypris (Fig. 5) 

Shells different; maxillule 3rd lobe with two Zahnborsten . . . . 14. 

14. Shell subtriangular, produced at each end; left valve overlapping right 
valve anteriorly and posteriorly, but overlapped by the right valve 
dorsally; maxillule palp distal segment spatulate Cyprilla (Fig. 5) 
Shell of a different type; maxillule palp elongate .. .. ..15. 

15. Shell small-medium; subtriangular to subovate in lateral view; ovate 
in dorsal view; left valve overlapping right valve anteriorly; anterior 
inner lamellae broad, posterior inner lamellae narrow . . . . 16. 
Shells of a different type . . . . . . . . . . . . . . 17. 

16. Shell typically whitish to greenish, usually with two or three irregular 
dark strigae on each valve in life; right valve denticulate anteriorly; 
inner lamellae as illustrated . . . . . . Cypridopsis (Fig. 4) 

Shell typically whitish; ornamented with pits arranged concentrically 

or else tuberculate; without strigae; inner lamellae as illustrated 

Zonocypris (Fig. 9) 

17. Shell small-medium to medium; subtriangular in lateral view; 
relatively compressed in dorsal view; right valve overlapping left valve 
anteriorly; anterior and posterior inner lamellae rather narrow; 
antennal natatory setae well developed . . . . . . . . . . 18. 

Shell small to small-medium; valves subequal; subreniform in lateral 
view; without strigae on the valves; anterior and posterior inner 
lamellae both relatively broad ; antennal natatory setae short 

Kap cypridopsis (Figs 6-7) 

18. In males, palp of first thoracic legs elongate, hemipenis anterior lobe 
curved upwards . . . . . . . . Plesiocypridopsis (Fig. 9) 

In males, palp of first thoracic legs of normal length, hemipenis 
anterior lobe pointing downwards . . Sarscypridopsis (Fig. 8) 

19. Shell small-medium to medium; smooth (Cyprettini, Bradycyprini) or 
pitted (Cyprettini) or striate (Bradycyprini); subglobose in lateral and 
dorsal view; radial septa well developed at least in right valve; furca 
'normal' but slender (Bradycyprini) or very slender (Cyprettini) 

Cyprettinae 20. 
Shell medium to very large, typically smooth; compressed to globose 
in dorsal view; without radial septa (exception Stenocypris, cf. 29); 
furca 'normal', not conspicuously slender . . . . . . . . 22. 

20. Radial septa developed in both valves . . CypTQttmi— Cypretta (Fig. 10) 
Radial septa developed in right valve only .. Bradycyprini 21. 

21. Shell striate Paracypretta (Figs 10-12) 

Shell smooth . . . . Bradycypris (Figs 11-12) 



52 ANNALS OF THE SOUTH AFRICAN MUSEUM 

22. Chitin supports with loculi where they branch proximally, furca 
flexuous . . . . . . . . . . . . Cypricercini 23. 

Chitin supports without loculi where they branch proximally; furca 
typically straight or evenly curved, rarely flexuous . . . . . . 25. 

23. Shell oval, with distinct posterior process in some species; chitin 
supports with anterior branch well developed; spermatic vesicles 

(J only) coiled anteriorly .. .. .. .. Cypricercus (Fig. 1 2) 

Shell oval-oblong, without posterior processes; chitin supports 
without an anterior branch . . . . . . . . . . . . 24. 

24. Furca normally developed (2 claws and 2 bristles) . . Tanycypris 
Furca with two claws only . . . . . . Gesa (cf. Introduction) 

25. Shell without a posterior inner lamella (the inner margin coincides 
posteriorly with the posterior shell margin cf. Rome 1965) . . . . 26. 
Shells with a posterior inner lamella . . . . . . . . . . 27. 

26. Distal segment of the P II with a short dorsal bristle (Rome 1965) 

Isocypridini, Isocypris (Fig. 13) 
Distal segment of the P II with a dorsal bristle which is almost as 
long as the terminal claw (Rome 1965) 

Amphibolocypridini, Amphibolocypris 

27. Chitin supports widened distally into triangular chitinous processes 

Herpetocypris, Stenocypridini 28. 
Chitin supports without distal triangular chitinous processes . . 31. 

28. Furcal rami similar, furca with a posterior bristle 

Herpetocypris (Figs 13-14) 
Furcal rami dissimilar; furca without a posterior bristle 

Stenocypridini 29. 

29. Shells with radial septa . . . . . . . . . . Stenocypris 

Shells without radial septa . . . . . . . . . . . . 30. 

30. Marginal pore canals very short . . . . . . . . Chrissia (Fig. 17) 

Marginal pore canals well developed and branched 

Parastenocypris (Figs 14-17) 

31. Either right or left valve of shell denticulate ventrally except 
Homocypris which, however, is of medium size only (length 1,1 -1,4 mm) 
and has a normal furca . . . . . . . . . . Cyprinotini 32. 

Neither right nor left valve of shell denticulate ventrally (except 
Apateleeypris cf. 42); carapace large, at least 1,75 mm, except 
Mesocypris (0,91 mm) which has an unique furca, cf. 34 . . . . 34. 

32. Shell elongate-subreniform in lateral view; anterior inner lamellae 
distinctly broader than posterior inner lamellae 

Homocypris (Figs 17-18) 
Shell elongate-ovate in lateral view; both anterior and posterior inner 
lamellae relatively, and subequally narrow . . . . . . . . 33. 

33. Right valve denticulate antero- and/or posteroventrally 

Heterocypris (Fig. 1 8) 



ILLUSTRATED GENERIC KEY TO SOUTH AFRICAN CONTINENTAL OSTRACODA 53 

Left valve denticulate antero- and/or posteroventrally Hemicypris 

34. Furca normally developed (2 claws, 2 bristles) and not modified as 
below . . . . . . . . . . . . . . . . . . 35. 

Furca with the posterior 'bristle 1 transformed into a stubby claw-like 
spine; furcal rami weakly dissimilar . . . . . . Mesocypris 

35. Carapace large, with prominent secondary fused zone in right valve 

Afrocypris (Figs 1 8-20) 
Carapace large, without prominent secondary fused zone in right valve 36. 

36. Carapace large; maxillule palp elongate .. .. .. ..37. 

Carapace large; maxillule palp relatively broad (i.e. approximately 
square or spatulate) . . . . . . . . . . . . . . 39. 

37. Carapace large, without ventral ala; penultimate 'segment' of P II 
divided . . . . . . . . . . Eucypridini, Eucypris (Fig. 19) 

Carapace large, with or without ventral ala; penultimate 'segment' of 

P II not divided . . . . . . . . . . . . Cypridini 38. 

38. Carapace large, with ventral ala . . . . . . Pseudocypris (Fig. 20) 

Carapace large, without ventral ala . . . . . . . . Cypris 

39. Carapace large, tuberculate in one species; antennal natatory setae 
well developed, reaching about as far as the distal part of the terminal 
segment . . . . . . . . . . . . Sclerocypris (Fig. 21) 

Carapace large, smooth to distinctly hirsute; antennal natatory setae 
strongly reduced . . . . . . . . . . . . . . . . 40. 

40. Carapace large, in general shape resembling Homocypris; PI palps 
(o* only) strikingly dissimilar on left and right limbs; hemipenes 
(o* only) unlike those of any other ostracode ; shape of female genital 
lobe also distinctive . . . . . . . . Liocypris (Fig. 22) 

Carapace large; general shape unlike that of Homocypris; PI palps 

(o* only) not strikingly dissimilar on left and right limbs, hemipenes 
(6* only) different to Liocypris . . . . . . Megalocypridini 41. 

41. Carapace large; with a small but distinct posteroventral caudal spine 

on each valve . . . . . . . . . . . . Hypselecypris 

Carapace large; without such a spine on either valve . . . . . . 42. 

42. Carapace large; in dorsal view indented at both ends; not denticulate 
posteroventrally . . . . . . . . Megalocypris (Figs 23-24) 

Carapace large; in dorsal view narrowing anteriorly, but rounded, not 
indented, posteriorly; denticulate posteroventrally Apatelecypris (Fig. 24) 

ACKNOWLEDGEMENTS 

I am grateful to the National Institute for Water Research, Pretoria and 
to the British Museum (Natural History) London for the opportunity to work 
in South Africa and to Dr R. G. Noble who organized my visit. My original 
drawings were traced for publication by Mr D. Goode of the Transvaal Museum. 
My thanks are due to Professor P. H. A. Sneath and Dr M. J. Sackin of the 



54 ANNALS OF THE SOUTH AFRICAN MUSEUM 

University of Leicester, who prepared the computer key of the appendix, using 
the PANKEY program. 

I am particularly grateful to Dr T. H. Barry, Director of the South African 
Museum, for the extended use of its laboratory and library facilities during my 
working visit to Cape Town, also to Dr B. F. Kensley at the South African 
Museum for his considerable assistance in reorganizing the illustrations. 

Mrs L. Vennell, Riverina College of Advanced Education, typed the 
manuscript. 

REFERENCES 

Hartmann, G. 1957. Ostracoden aus dem Namaland und Transvaal. Veroff. naturw. 

Osnabruck 28: 50-60. 
McKenzie, K. G. \91\a. Species list of South African freshwater Ostracoda with an appendix 

listing museum collections and some further determinations. Ann. S. Afr. Mus. 57: 

157-213. 
McKenzie, K. G. \91\b. Ostracoda from Lake Peunde, near Mt. Wilhelm, New Guinea. 

Zool. Anz. 186: 391-403. 
McKenzie, K. G. 1972. Results of the speleological survey in South Korea 1966. XXII. 

Subterranean Ostracoda from South Korea. Bull. Natl. Sci. Mus., Tokyo 15: 155-166. 
Moore, R. C. ed. 1961. Treatise on Invertebrate Paleontology. Part Q. Arthropoda 3 Crustacea 

Ostracoda. Lawrence: Geological Society of America and University of Kansas Press. 
Rome, D. R. 1965. Crustacea: Ostracoda. In Hanstrom, B., Brinck, P. & Rudebeck, G., eds. 

South African animal life 11 : 9-58. Stockholm : Swedish Natural Science Research Council. 
Sars, G. O. 1896. On some South African Entomostraca raised from dried mud. Skr. 

VidenskSelsk. Christiania 1895 (8): 1-56. 
Sars, G. O. 1924a. The freshwater Entomostraca of the Cape Province (Union of South 

Africa). Part II: Ostracoda. Ann. S. Afr. Mus. 20: 105-193. 
Sars, G. O. 19246. Contributions to a knowledge of the fauna of South West Africa. 

Ann. S. Afr. Mus. 20: 195-211. 



APPENDIX 

South African Freshwater Ostracoda Generic Key 

(prepared by the Program PANKEY). 

1 . Shell : surface not smooth 2. 

2. Furca reduced to whip-like seta 3. 

3. Dorsal view of shell broad 4. 

4. Maxillule 3rd lobe with 4 or more Zahnborsten, maxilla epipod with 

less than 3 Strahlen, shell denticulation in one or both valves absent, 
shell: radial septa or lunettes in inner lamellae present in both valves, 
shell surface pustulose not tuberculate-costate. Oncocypris 

4. Maxillule 3rd lobe with 2 Zahnborsten, maxilla epipod with more than 

3 up to 6 Strahlen, shell denticulation in one or both valves present, 
shell : radial septa or lunettes in inner lamellae, shell surface tuberculate- 
costate not pustulose Zonocypris 
3. Dorsal view of shell 'normal' 5. 
5. Parthenogenesis: males virtually unknown, shell denticulation in one, 

or both valves present. Cypridopsis 

5. Parthenogenesis : both sexes known 6. 

6. Maxilla male epipod not elongated. Cyprilla 

6. Maxilla male epipod elongated Plesiocypridopsis 



ILLUSTRATED GENERIC KEY TO SOUTH AFRICAN CONTINENTAL OSTRACODA 55 

2. Furca 'normal' 7. 

7. 7th limb reflexed, terminal seta long 8. 

8. Maxilla endopod 2-segmented, chitin supports: distally with 

simple point, Zenker's Organ with more than 15 whorls, shell: 
dorsal sulci present Ilyocypris 

8. Maxilla endopod unisegmented 9. 

9. Shell denticulation in one or both valves present Physocypria 

9. Shell denticulation in one or both valves absent Cyclocypris 

7. 7th limb reflexed, terminal seta short 10. 

10. Furcal rami very slender, Zenker's Organ with 7—14 whorls, 

shell : radial septa or lunettes in inner lamellae present in both 
valves, shell length 0,45-0,90 mm Cypretta 

10. Zenker's Organ with more than 15 whorls 1 1. 

11. Furcal rami slender, shell: radial septa or lunettes in inner 

lamellae present in one valve, shell surface striate, shell length 
0,90-1 ,75 mm Paracypretta 

11. Furcal rami 'normal' 12. 

12. Dorsal view of shell broad Cypris 

12. Dorsal view of shell 'normal' Sclerocypris 

1. Shell : surface smooth (includes weak pitting) 13. 

13. 7th limb penultimate segment undivided 14. 

14. Parthenogenesis: both sexes known, maxilla epipod with less than 

3 Strahlen Candonopsis 

14. Parthenogenesis: males virtually unknown 15. 

15. Mandible palp comb present, maxilla epipiod with more than 

7 Strahlen, maxilla endopod 3-segmented, 7th limb not reflexed, 
furca absent, shell : inner lamellae absent. Darwinula 

15. Mandible palp comb absent, maxilla epipod with more than 3 up to 

6 Strahlen, maxilla endopod unisegmented (but 'ghost' segmentation 
indicated), 7th limb reflexed, furca 'normal', shell: inner lamellae 
present. Paracypris 

13. 7th limb penultimate segment divided 16. 

16. Natatory setae of antennae weakly developed 17. 

17. Maxilla epipod with less than 3 Strahlen, furca: posterior margins 

dissimilar, shell densely hirsute Mesocypris 

17. Maxilla epipod with more than 3 up to 6 Strahlen 18. 

18. Furca reduced to whip-like seta, Zenker's Organ with 7-14 whorls, 

shell length 0,45-0,90 mm Kapcypridopsis 

18. Furca 'normal' 19. 

19. Chitin supports: distally with simple point Liocypris 

19. Chitin supports : distally elaborated 20. 

20. Shell denticulation in one or both valves absent, shell length 

3,60-7,30 mm Megalocypris 

20. Shell denticulation in one or both valves present, shell length 

1,75-3,60 mm Apatelecypris 

16. Natatory setae of antennae well developed 21. 

21. Chitin supports : distally elaborated 22. 

22. Furca: posterior bristle absent 23. 

23. Shell length 1,75-3,60 mm 24. 

24. Shell: radial septa or lunettes in inner lamellae present in 

both valves Stenocypris 

24. Shell: radial septa or lunettes in inner lamellae absent 

'Parastenocypris' 

23. Shell length 0,90-1,75 mm 25. 

25. 6th limb: distal bristle almost as long as the distal claw, 

furca: posterior margins similar, chitin supports: 

triangular process absent, dorsal view of shell very 

compressed, shell: inner lamellae absent posteriorly 

Amphibolocypris 



56 ANNALS OF THE SOUTH AFRICAN MUSEUM 

25. 6th limb: distal bristle short, furca: posterior margins 

dissimilar, chitin supports: triangular process present, 
dorsal view of shell normal, shell : inner lamellae present Chrissia 
22. Furca : posterior bristle present 26. 

26. Chitin supports: triangular process present, shell length 

1,75-3,60 mm Herpetocypris 

26. Chitin supports: triangular process absent 27. 

27. Dorsal view of shell very compressed, shell: inner 

lamellae absent posteriorly, shell length 0,90-1,75 mm Isocypris 
27. Dorsal view of shell 'normal' 28. 

28. Shell posterior processes absent Afrocypris 

28. Shell posterior processes present Hypselecypris 

21. Chitin supports : distally with simple point 29. 

29. Dorsal view of shell broad 30. 

30. Furcal rami slender, shell : radial septa or lunettes in 

inner lamellae present in one valve, shell length 
0,90-1,75 mm Bradycypris 

30. Furcal rami 'normal' 31. 

31. Dorsal view of shell: alae absent Eucypris 

3 1 . Dorsal view of shell : alae present Pseudocypris 

29. Dorsal view of shell 'normal' 32. 

32. Chitin supports: loculi present where they branch 

proximally 33. 

33. Chitin supports: loculi present and prominent 

anterior branch, shell posterior processes present 

Cypricercus 
33. Chitin supports: loculi present but without 

prominent anterior branch, shell posterior 
processes absent Tanycypris 

32. Chitin supports: loculi absent where they branch 

proximally 34. 

34. Shell denticulation in one or both valves absent, 

shell inner lamellae: anterior broad posterior 
narrow Homocypris 

34. Shell denticulation in one or both valves present 35. 

35. Shell denticulation in right valve Heterocypris 

35. Shell denticulation in left valve Hemicypris 

REFERENCE 

Pankhurst, R. J., 1970. A computer program for generating diagnostic keys. Computer J. 12: 
145-151. 



Fig. 1 

1. Darmnula sp.; ovigerous ?; adductor muscle scars, left valve, x 175. 

2. Darwinula sp.; ovigerous ?; outline right valve, x 125. 

3. Generalized rimmed normal pore canal, x 1400. 

4. Generalized rimless normal pore canal, x 2500. 

5. Generalized sieve type normal pore canal, x 1000. 

6. Generalized lophodont hingement right valve (after Moore ed. 1961). 

7. Generalized merodont hingement right valve (after Moore ed. 1961). 

8. Generalized entomodont hingement right valve (after Moore ed. 1961). 

9. Generalized gonglyodont hingement right valve (after Moore ed. 1961). 

10. Cyprideis sp. ; hingement right valve, x 100. 

11. Cyprideis sp.; adductor muscle scars and fulcral scar (dashes), x 225. 

12. Loxoconcha sp.; adductor muscle scars and fulcral scar (dashes), x 300. 

13. Loxoconcha sp.; anterior hingement right valve, x 250. 



ILLUSTRATED GENERIC KEY TO SOUTH AFRICAN CONTINENTAL OSTRACODA 57 



1 
3 

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8 



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c? 



CO 



<£> 



11 




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'3 o 

so 



RD 




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13 



58 



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ILLUSTRATED GENERIC KEY TO SOUTH AFRICAN CONTINENTAL OSTRACODA 59 



Fig. 2 

1. Gomphocythere expansa Sars, 1924; SAM-A11303; ovigerous $; P I and P II. x 400. 

2. Posterior of body with caudal lobes, x 400. 

3. P III. X 400. 

4. Antennule. x 400. 

5. Antenna, x 400. 






60 



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ILLUSTRATED GENERIC KEY TO SOUTH AFRICAN CONTINENTAL OSTRACODA 61 



Fig. 3 

1. Ilyocypris australiensis Sars, 1889; SAM-A11; adult ?; detail anterior margin right valve 
showing radial pore canals and muscle scars, x 160. 

2. Antennule, six terminal segments, x 400. 

3. P III, terminal segments, x 400. 

4. PI. x 400. 

5. Mandible palp, x 400. 

6. Furca. x 400. 

7. Physocypria capensis (Sars, 1896); SAM-A11136; adult S; internal view right valve. 

x 160. 



62 



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Fig. 4 

1. Physocypria capensis (Sars, 1896); SAM-A11136; adult <J; Zenkers Organ, x 400. 

2. Mandible palp, detail j8 and y bristles, x 400. 

3. Antennule, six terminal segments, x 400. 

4. P III, terminal segment, x 400. 

5. Cypridopsis viduella Sars, 1896; SAM-A11171 ; adult ?; P I. x 400. 

6. Rake-like organ. X 400. 

7. P III, terminal segment, x 400. 

8. Antennule, six terminal segments, x 400. 

9. Mandible palp, detail a, j8 and y bristles, x 400. 
10. Internal view right valve, x 160. 



64 



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ILLUSTRATED GENERIC KEY TO SOUTH AFRICAN CONTINENTAL OSTRACODA 65 



Fig. 5 

1. Oncocypris cf. voeltzkowi Miiller, 1898; SAM-A11374; ovigerous ?; maxillule palp and 
lobes, third lobe with some Zahnborsten. x 400. 

2. PI. x 400. 

3. Anterior margin of left valve, x 160. 

4. Mandible palp, detail of a, /3 and y bristles, x 400. 

5. Antennule, six terminal segments, x 400. 

6. Cyprilla arcuata Sars, 1 924; SAM-A1 1 1 58 ; ovigerous 9 ; antennule, six terminal segments. 
x 400. 

7. Internal view left valve, x 160. 

8. Maxillule palp. X 400. 

9. P II. x 400. 

10. Antenna, first endopod segment. K 400. 

1 1 . P III. x 400. 



66 



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Fig. 6 

1. Kapcypridopsis barnardi McKenzie, 1977; SAM-A15451; adult $ (yellowish form); 
internal view left valve, x 160. 

2. Antennule segments, x 400. 

3. Antenna, with 'natatory' setae, x 400. 

4. Rake-like organ, x 400. 

5. Mandible palp, detail of a, |8 and y bristles, x 400. 

6. Maxillule palp and lobes, x 400. 

7. P III, terminal segments, x 400. 

8. P II. x 400. 

9. Furca. x 400. 
10. PI. x 400. 



68 



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ILLUSTRATED GENERIC KEY TO SOUTH AFRICAN CONTINENTAL OSTRACODA 69 



Fig. 7 

1. Kapcypridopsis bamardi McKenzie, 1977; SAM-A15451 ; adult <$ (greenish form); internal 
view right valve, x 160. 

2. PI right palp, x 400. 

3. P II. X 400. 

4. Zenkers Organ, x 400. 

5. Eye. x 400. 

6. Antennule, six terminal segments, x 400. 

7. Hemipenis. x 400. 



70 



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Fig. 8 

1. Sarscypridopsis striolata (Sars, 1924); SAM-A 11201; ovigerous ?; internal view right 
valve, x 160. 

2. Sarscypridopsis ochracea (Sars, 1924); SAM-A1 1220; adult S\ mandible palp detail 
a, j8 and y bristles, x 400. 

3. Antennule, six terminal segments, x 400. 

4. Antenna, first endopod segment with sensory bristle, x 400. 

5. Posteroventral right valve, x 160. 

6. P II. x 400. 

7. Hemipenis. x 400. 

8. Maxillule third lobe with Zahnborsten. x 400. 

9. Muscle scars, x 160. 

10. PI, left palp, x 400. 

11. P III, terminal segment, x 400. 

12. Maxillule palp, x 400. 

13. PI, right palp, x 400. 



72 



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Fig. 9 

1. Plesiocypridopsis cf. aldabrae (Muller, 1898); SAM-A5940; adult £; P 1 right palp. 

x 400. 

2. P I left palp, x 400. 

3. Antenna, sensory bristle first endopod segment, x 400. 

4. Zonocypris cordata Sars, 1924; SAM-A11 144; ovigerous $; anterior margin left valve. 

x 160. 

5. Plesiocypridopsis cf. aldabrae (Muller, 1898); SAM-A5940; adult ?; ventral view of labia 
with rake-like organs, x 400. 

6. Mandible palp, detail of a, ft and y bristles, x 400. 

7. Antennule, six terminal segments, x 400. 

8. P III, terminal segment, x 400. 

9. Adult c?; head and oral region, x 160. 

10. Anteroventral right valve, x 160. 

11. Hemipenis. x 400. 



74 



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ILLUSTRATED GENERIC KEY TO SOUTH AFRICAN CONTINENTAL OSTRACODA 75 



Fig. 10 

1. Cypretta cf. turgida (Sars, 1895); SAM-A1 1 159; ovigerous ?; anterior margin right valve 
with radial septa, x 160. 

2. Antenna endopod segments with sensory bristle. x 400. 

3. Antennule six terminal segments, x 400. 

4. Mandible palp, detail of a, j3 and y bristles, x 400. 

5. Maxillule palp and third lobe with Zahnborsten. x 400. 

6. P I, terminal segments, x 400. 

7. Distal furca. x 400. 

8. Paracypretta ampullacea Sars, 1924; SAM-A11278; ovigerous $; sensory bristle first 
endopod segment of antenna, x 400. 

9. P III, distal terminal segment, x 400. 

10. Mandible palp, detail of a, /3 and y bristles, x 400. 

11. Maxillule palp and third lobe with Zahnborsten. x 400. 



76 



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Fig. 11 

1. Paracypretto ampullacea Sars, 1924; SAM-A1 1278; ovigerous 9; internal view left valve. 

x 40. 

2. Chitin support, x 160. 

3. Detail anterior margin radial pore canals left valve, x 160. 

4. Bradycypris intumescens (Brady, 1907); SAM-A11299; ovigerous $; internal view right 
valve, showing radial septa, x 40. 

5. Rake-like organ, x 400. 

6. Antennule, six terminal segments, x 160. 

7. Antenna, sensory bristle first endopod segment, x 400. 

8. Maxillule palp and lobes, third lobe with Zahnborsten. x 160. 

9. Mandible palp, detail of a, /3 and y bristles, x 400. 
10. Mandible coxale. x 160. 



78 



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Fig. 12 

1. Paracypretta ampullaeea Sars, 1924; SAM-A1 1278; ovigerous ?; furca. x 160. 

2. Eye. x 400. 

3. Antennule, terminal six segments, x 160. 

4. Antennule, detail second segment, x 400. 

5. Bradycypris intumescens (Brady, 1907); SAM-A 11299; ovigerous $; furca. x 160. 

6. Chitin support, x 160. 

7. Cypricercus cuneatus Sars, 1896; SAM-A 1 1 140; adult <$ ; antennule, detail first and second 
segments with gland and chemo-receptor. x 400. 

8. Chitin support, x 160. 

9. Antenna, sensory bristle of first endopod segment, x 400. 

10. Cap of Zenkers Organ, x 400. 

1 1 . Mandible palp, detail of a, j3 and y bristles, x 400. 

12. PI, right palp, x 400. 

13. Furca. x 160. 

14. PI, left palp, x 400. 



80 



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Fig. 13 

1. Isocypris priomena Miiller, 1908; SAM-A11177; ovigerous ?; antennule second segment. 
x 400. 

2. Internal view left valve, x 40. 

3. P II, three terminal segments, x 160. 

4. Antenna endopod segments, x 160. 

5. Anterior margin, detail radial pore canals, x 160. 

6. Mandible palp, detail of a, j3 and y bristles, x 160. 

7. P III terminal segment, x 400. 

8. Maxillule palp, x 160. 

9. Maxillule third lobe with Zahnborsten. x 160. 

10. Herpetocypris chevreuxi (Sars, 1896); SAM-A1148; ovigerous ?; antennule second and 
third segments, x 160. 

11. Antennule second segment chemo-receptor. x 400. 

12. Rake-like organ, x 160. 

13. P III, terminal segment, x 160. 

14. Internal view left valve, x 40. 



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Fig. 14 

1. Herpetocypris chevreuxi (Sars, 1896); SAM-A1148; ovigerous ?; maxillule third lobe. 
x 400. 

2. Chitin support, x 160. 

3. P I, terminal segments and part of claw, x 160. 

4. Mandible palp, detail of a, £ and y bristles, x 400. 

5. Parastenocypris declivis (Sars, 1924); SAM-A3797; ovigerous $; internal view, left valve. 

x 40. 

6. Distal furca. x 160. 

7. Mandible palp, detail of a, /3 and y bristles, x 160. 

8. Antennule second segment with chemo-receptor. x 400. 



84 



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Fig. 15 

1. Parastenocypris declivis (Sars, 1924); SAM-A3797; adult ?; chitin support. X 160. 

2. Detail anterior margin left valve, x 160. 

3. Detail posteroventral margin left valve, x 160. 

4. Maxillule third lobe with Zahnborsten. x 400. 

5. Parastenocypris hodgsoni (Sars, 1924); SAM-A11126; ovigerous $; internal view left 
valve, x 16. 

6. Furca, detail of posterior margin, x 160. 

7. Detail anteroventral margin left valve, x 160. 

8. Furca. x 40. 

9. Furca, other ramus, x 160. 

10. Antennule, detail of second segment chemo-receptor. x 400. 

11. Parastenocypris pardalis (Sars, 1924); SAM-A11167; internal view right valve, x 16. 

12. Internal view left valve, x 16. 

13. Antenna, sensory bristle of basal endopod segment, x 400. 



86 



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87 



Fig. 16 

1. Parastenocypris partialis (Sars, 1924); SAM-A1 1 167; adult j ; detail anterior margin left 
valve, x 160. 

2. PI, left palp, x 160. 

3. PI, right palp, x 160. 

4. Antennule second and third segments, x 160. 

5. Parastenocypris olivacea (Sars, 1924); SAM-A3797; adult <$\ internal view left valve. 

X 16. 

6. Muscle scars, x 160. 

7. Detail anterior margin left valve, x 160. 

8. Antenna terminal segment, detail of comb bristle, x 400. 

9. Mandible palp, detail of a and j8 bristles, x 400. 

10. Mandible palp, detail of y bristle, x 400. 

11. Terminal rake-like organ, x 160. 

12. Antenna basal endopod segment, showing exopod and sensory bristle, x 400. 

13. Hem i pen is. X 160. 



88 



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Fig. 17 

1. Parastenocypris olivacea (Sars, 1924); SAM-3793; adult 3; distal furca. x 160. 

2. Frontal head region and labrum. x 160. 

3. IChrissia fascigera (Sars, 1924); SAM-A11361 ; adult ?; internal view posteroventral left 
valve, x 160. 

4. Anteroventral margin left valve, showing simple nature of radial pore canals, x 400. 

5. Homocypris conoidea Sars, 1924; SAM-A11170; adult $; detail anterior margin right 
valve, x 160. 

6. Muscle scars and detail of shell microstructure. x 160. 

7. P III, terminal segments, x 160. 

8. Chitin support, x 160. 

9. Internal view right valve, x 40. 

10. Detail anteroventral margin right valve, x 160. 

11. Furca. x 160. 

12. Antenna basal endopod segment, showing exopod, sensory bristle and part of the 'natatory' 
setae, x 160. 



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Fig. 18 

1. Homocypris conoidea Sars, 1924; SAM-AI 1 170; adult ?; mandible palp, detail of a, /3 
and y bristles, x 400. 

2. Rake-like organs, x 400. 

3. Antennule, detail second segment, x 400. 

4. Maxillule third lobe with Zahnborsten. x 400. 

5. P I, endopod and exopod, several bristles not shown, x 160. 

6. Heterocypris aurea (Sars, 1896); SAM-AI 1310; ovigerous $; antennule, terminal six 
segments, x 160. 

7. Mandible palp, detail of a, /3 and y bristles, x 160. 

8. P III, terminal segment, x 160. 

9. Internal view right valve, x 40. 

10. Maxillule palp and lobes, third lobe with Zahnborsten. x 160. 

11. Furca. x 160. 

12. P II, four terminal segments, x 160. 

13. Afrocypris barnardi Sars, 1924; SAM-AI 1291; ovigerous ?; maxillule palp, second 
segment, x 160. 

14. Detail, anterior margin right valve, x 40. 

15. Internal view right valve, x 16. 



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Fig. 19 

1. Afrocypris barnardi Sars, 1924; SAM-A11291 ; ovigerous $; antennule, detail of second 
segment, x 160. 

2. Antenna, endopod segments, x 40. 

3. Antenna, detail of sensory bristle on basal endopod segment, x 160. 

4. Rake-like organ, x 160. 

5. Mandible palp, detail of a, j3 and y bristles, x 160. 

6. P III, terminal segment, x 40. 

7. Distal furca. x 40. 

8. Antennule, five terminal segments, x 40. 

9. Chitin support, x 40. 

10. P II, three terminal segments, x 40. 

11. Eucypris purcelli Sars, 1924; SAM-A11287; adult ?; P II. x 160. 

12. Distal furca. x 160. 

13. Maxillule palp and third lobe showing Zahnborsten. x 160. 

14. PI, exopod bristles not shown, x 160. 

15. Antennule, detail of chemo-receptor. x 400. 

16. Antennule, six terminal segments, x 160. 



94 



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Fig. 20 

1. Afrocypris barnardi Sars, 1924: SAM A 1 1291; ovigerous _; maxillule third lobe, x 160. 

2. Pseudocypris triquetra Sars, 1924; SAM-A11211; adult 5; maxillule palp and lobes. 

x 160. 

3. PI, right palp, x 160. 

4. P 1, left palp, x 160. 

5. P II, two terminal segments and claw. :: 160. 

6. Outline hemipenes. x 40. 

7. Chitin support, x 160. 

8. Distal furca. x 160. 

9. Antenna, sensory bristle, x 160. 

10. P III, distal segment, x 160. 

11. Antennule, six distalmost segments, x 160. 

12. Maxillule third lobe showing Zahnborsten. > 160. 

13. Rake-like organ, x 160. 

14. Mandible palp, detail a, /3 and y bristles, x 160. 



96 



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Fig. 21 

1. Sclerocypris clavularis Sars, 1924; SAM-A1 11 18; ovigerous 9; antennule, six distalmost 
segments, x 160. 

2. Rake-like organ, x 160. 

3. PI, four terminal segments, x 160. 

4. Chitin support, x 160. 

5. Maxillule palp and third lobe showing Zahnborsten. x 160. 

6. Carapace, muscle scars, x 40. 

7. Mandible palp, detail a, /? and y bristles, x 160. 

8. Anterior margin left valve, x 160. 

9. Anterior margin right valve, x 160. 

10. Internal view right valve, x 16. 

11. Distal furca. x 160. 



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Fig. 22 

1. Liocypris grandis Sars, 1924; SAM-6281; adult $; carapace, detail anterior margin 
reticulate valve microstructure (in part), x 40. 

2. Internal view right valve, x 16. 

3. Furca, distal detail, x 160. 

4. Chitin support, x 40. 

5. Antennule, detail of second segment showing gland and bristle. x 160. 

6. Maxillule third lobe with Zahnborsten. x 160. 

7. Maxillule palp, x 160. 

8. P I palp, x 40. 

9. Rake-like organ, x 160. 

10. Mandible palp, indicating complexity of setation. x 160. 

11. Mandible epipod. x 160. 



100 



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Fig. 23 

1. Megalocypris hispida Sars, 1924; SAM-A11845; adult $; mandible palp, detail of a, /3 
and y bristles. X 160. 

2. SAM-A11844; adult S\ antennule, six distalmost segments, x 40. 

3. SAM-A11845; adult $; mandible coxale, detail of toothbrush bristles between first and 
second teeth, x 400. 

4. Megalocypris sp. (smooth form with hemipenis similar to M. tuberculata Sars, 1924); 
SAM-A11849; adult cT; antenna, endopod segments, x 40. 

5. and 6. Rake-like organs, x 160. 

7. Mandible palp, detail of a and /3 bristles. Note that this male carries only one y bristle. 
x 160. 

8. Megalocypris princeps Sars, 1898; SAM-1479; adult <$; rake-like organ, x 160. 

9. and 10. Megalocypris tuberculata Sars, 1924; SAM-A11273; adult <$; rake-like organs. 

x 160. 

11. PI palp, x 40. 

12. Furca. X 40. 

13. Chitin support, x 40. 

14. P II, three terminal segments, x 160. 

15. Maxillule palp, x 160. 

16. Maxillule third lobe with Zahnborsten. x 160. 

17. Antenna, sensory bristle of first endopod segment, x 160. 

18. Carapace, muscle scars, x 40. 

19. Outline of hemipenis. x 40. 



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Fig. 24 

1. Megalocypris tuberculata Sars, 1924; SAM-A11273; adult #; mandible palp, detail of a, 
j3 and y bristles. Note that the specimen may be a freak since it carries two y bristles 
instead of the normal complement of one only, x 160. 

2. Antennule, detail of second segment showing gland, chemo-receptor and bristle, x 160. 

3. Apatelecypris brevis Sars, 1924; SAM-A11369; adult <$; carapace, internal view right 
valve, muscle scars not shown, x 16. 

4. Detail anteroventral radial pore canals, x 160. 

5. Detail anterior margin right valve, x 40. 

6. Antenna, detail 'natatory' setae, x 160. 

7. PI, detail of left palp, x 160. 

8. P I, detail of right palp, x 160. 

9. Antennule, detail of second segment showing chemo-receptor and bristle, x 160. 

10. Mandible palp, detail of y bristle, x 160. 

1 1. Mandible palp, detail of a and /3 bristles, x 160. 

12. Tips of rake-like organs, x 160. 

13. Furca. x 40. 

14. Maxillule, palp and third lobe showing Zahnborsten. x 160. 

15. Chitin support, x 40. 

16. P II, three terminal segments, x 160. 



6. SYSTEMATIC papers must conform with the International code of zoological nomenclature 
(particularly Articles 22 and 51). 

Names of new taxa, combinations, synonyms, etc., when used for the first time, must be 
followed by the appropriate Latin (not English) abbreviation, e.g. gen. nov., sp. nov., comb, 
nov., syn. nov., etc. 

An author's name when cited must follow the name of the taxon without intervening 
punctuation and not be abbreviated; if the year is added, a comma must separate author's 
name and year. The author's name (and date, if cited) must be placed in parentheses if a 
species or subspecies is transferred from its original genus. The name of a subsequent user of 
a scientific name must be separated from the scientific name by a colon. 

Synonymy arrangement should be according to chronology of names, i.e. all published 
scientific names by which the species previously has been designated are listed in chronological 
order, with all references to that name following in chronological order, e.g. : 

Family Nuculanidae 

Nuculana (Lembulus) bicuspidata (Gould, 1845) 

Figs 14-1 5A 
Nucula (Leda) bicuspidata Gould, 1845: 37. 
Leda plicifera A. Adams, 1856: 50. 

Laeda bicuspidata Hanley, 1859: 118, pi. 228 (fig. 73). Sowerby, 1871 : pi. 2 (figs 8a-b). 
Nucula largillierti Philippi, 1861 : 87. 
Leda bicuspidata: Nickles, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9. 

Note punctuation in the above example : 

comma separates author's name and year 

semicolon separates more than one reference by the same author 

full stop separates references by different authors 

figures of plates are enclosed in parentheses to distinguish them from text-figures 

dash, not comma, separates consecutive numbers 

Synonymy arrangement according to chronology of bibliographic references, whereby 
the year is placed in front of each entry, and the synonym repeated in full for each entry, is 
not acceptable. 

In describing new species, one specimen must be designated as the holotype; other speci- 
mens mentioned in the original description are to be designated paratypes ; additional material 
not regarded as paratypes should be listed separately. The complete data (registration number, 
depository, description of specimen, locality, collector, date) of the holotype and paratypes 
must be recorded, e.g. : 

Holotype 

SAM-A13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King's Beach, 
Port Elizabeth (33°51'S 25°39'E), collected by A. Smith, 15 January 1973. 

Note standard form of writing South African Museum registration numbers and date. 

7. SPECIAL HOUSE RULES 

Capital initial letters 

(a) The Figures, Maps and Tables of the paper when referred to in the text 

e.g. '. . . the Figure depicting C. namacolus . . .'; *. . . in C. namacolus (Fig. 10) . . .' 

(b) The prefixes of prefixed surnames in all languages, when used in the text, if not preceded 
by initials or full names 

e.g. Du Toit but A. L. du Toit; Von Huene but F. von Huene 

(c) Scientific names, but not their vernacular derivatives 
e.g. Therocephalia, but therocephalian 

Punctuation should be loose, omitting all not strictly necessary 
Reference to the author should be expressed in the third person 
Roman numerals should be converted to arabic, except when forming part of the title of a 

book or article, such as 

'Revision of the Crustacea. Part VIII. The Amphipoda.' 
Specific name must not stand alone, but be preceded by the generic name or its abbreviation 

to initial capital letter, provided the same generic name is used consecutively. 
Name of new genus or species is not to be included in the title : it should be included in the 

abstract, counter to Recommendation 23 of the Code, to meet the requirements of 

Biological Abstracts. 



K. G. McKENZIE 

ILLUSTRATED GENERIC KEY TO 
SOUTH AFRICAN CONTINENTAL OSTRACODA 



VOLUME 74 PART 4 NOVEMBER 1977 



ISSN 0303-2515 



MUS. COMP. ZOOU 
LIBRARY 

'JAN 3 1 1978 

HARVARD 
UNIVERSITY 



ANNALS 




INSTRUCTIONS TO AUTHORS 

1. MATERIAL should be original and not published elsewhere, in whole or in part. 

2. LAYOUT should be as follows: 

(a) Centred masthead to consist of 

Title : informative but concise, without abbreviations and not including the names of new genera or species 

Author's(s') name(s) 

Address(es) of author(s) (institution where work was carried out) 

Number of illustrations (figures, enumerated maps and tables, in this order) 

(b) Abstract of not more than 200 words, intelligible to the reader without reference to the text 

(c) Table of contents giving hierarchy of headings and subheadings 

(d) Introduction 

(e) Subject-matter of the paper, divided into sections to correspond with those given in table of contents 

(f ) Summary, if paper is lengthy 

(g) Acknowledgements 
(h) References 

(i) Abbreviations, where these are numerous 

3. MANUSCRIPT, to be submitted in triplicate, should be typewritten and neat, double spaced 
with 2,5 cm margins all round. First lines of paragraphs should be indented. Tables and a list of 
legends for illustrations should be typed separately, their positions indicated in the text. All 
pages should be numbered consecutively. 

Major headings of the paper are centred capitals; first subheadings are shouldered small 
capitals; second subheadings are shouldered italics; third subheadings are indented, shouldered 
italics. Further subdivisions should be avoided, as also enumeration (never roman numerals) 
of headings and abbreviations. 

Footnotes should be avoided unless they are short and essential. 

Only generic and specific names should be underlined to indicate italics; all other marking 
up should be left to editor and publisher. 

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All illustrations, whether line drawings or photographs, should be termed figures (plates 
are not printed; half-tones will appear in their proper place in the text) and numbered in a 
single series. Items of composite figures should be designated by capital letters; lettering of 
figures is not set in type and should be in lower-case letters. 

The number of the figure should be lightly marked in pencil on the back of each illustration. 

5. REFERENCES cited in text and synonymies should all be included in the list at the end of 
the paper, using the Harvard System (ibid., idem, loc. cit., op. cit. are not acceptable): 

(a) Author's name and year of publication given in text, e.g. : 

'Smith (1969) describes . . .' 

'Smith (1969: 36, fig. 16) describes . . .' 

'As described (Smith 1969a, 19696; Jones 1971)' 

'As described (Haughton & Broom 1927) . . .' 

'As described (Haughton et al. 1927) . . .' 

Note: no comma separating name and year 

pagination indicated by colon, not p. 

names of joint authors connected by ampersand 

et al. in text for more than two joint authors, but names of all authors given in list of references. 

(b) Full references at the end of the paper, arranged alphabetically by names, chronologically 
within each name, with suffixes a, b, etc. to the year for more than one paper by the same 
author in that year, e.g. Smith (1969a, 19696) and not Smith (1969, 1969a). 

For books give title in italics, edition, volume number, place of publication, publisher. 

For journal article give title of article, title of journal in italics (abbreviated according to the World list o 
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number (only if independently paged) in parentheses, pagination (first and last pages of article). 

Examples (note capitalization and punctuation) 

Bullough, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan. 

Fischer, P.-H. 1948. Donnees sur la resistance et de le vitalite des mollusques. /. Conch., Paris 88: 100-140. 

Fischer, P.-H., Duval, M. & Raffy, A. 1933. Etudes sur les echanges respiratoires des littorines. Archs 

Zool. exp. gen. 74: 627-634. 
Kohn, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon. 

Ann. Mag. nat. Hist. (13) 2: 309-320. 
Kohn, A. J. 19606. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean. 

Bull. Bingham oceanogr. Coll. 17 (4): 1-51. 
Thiele, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. In: Schtjltze, L. Zoologische 

und anthropologische Ergebnisse einer Forschungsreise im west lichen und zentralen Sud-Afrika 4 : 269-270. 

Jena: Fischer. Denkschr. med.-naturw. Ges. Jena 16: 269-270. 

(continued inside back cover) 



ANNALS OF THE SOUTH AFRICAN MUSEUM 
ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM 



Volume 74 Band 
November 1977 November 
Part 4 Deel 




THE SOUTH AFRICAN MUSEUM'S 

MEIRING NAUDE CRUISES 

PART 6 

AMPHIPODA 



By 
CHARLES GRIFFITHS 



Cape Town Kaapstad 



The ANNALS OF THE SOUTH AFRICAN MUSEUM 

are issued in parts at irregular intervals as material 
becomes available 

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THE SOUTH AFRICAN MUSEUM'S MEIRING NAUDE CRUISES 

PART 6 
AMPHIPODA 

By 

Charles Griffiths 
Zoology Department, University of Cape Town 

(With 10 figures) 
[MS. accepted 4 August 1977] 

ABSTRACT 

Amphipoda recovered during the 1975, 1976 Meiring Naude cruises are listed and species 
of note discussed. The collection includes two new genera— Izinkala gen. nov. and Valettiella 
gen. nov. (Lysianassidae) — represented by /. fihla sp. nov. and V. castellana sp. nov. Three 
further species are recorded from southern Africa for the first time and Bathyamaryllis 
conocephala (K. H. Barnard) is redescribed. 



CONTENTS 



Introduction 
List of species . 
Systematic discussion 
Acknowledgements 
References 



PAGE 

105 
106 
108 
123 
123 



INTRODUCTION 

The material described herein is derived from collections made by the staff 
of the South African Museum during exploratory cruises by the R.V. Meiring 
Naude in 1975 and 1976. These cruises are of particular interest since they have 
investigated deeper waters off Natal which have previously received scant 
attention, despite their potential zoogeographical interest. A full listing of 
station data from both cruises has been presented elsewhere (Louw 1977), but 
few of these samples in fact used gear suitable for the capture of benthic 
amphipods. Although the number of amphipods recovered was limited, the 
proportion of new or unusual species in the collection bears witness to the 
potential for further exploration of this area. 

All material described below has been deposited in the South African 
Museum, Cape Town. 

105 
Ann. S. Afr. Mus. 74 (4), 1977: 105-123, 10 figs. 



106 ANNALS OF THE SOUTH AFRICAN MUSEUM 

LIST OF SPECIES 

SM Station Number of 
Number Individuals 

Suborder Gammaridea 

Family Acanthonotozomatidae 

Dikwa acrania Griffiths 86 2 

Family Ampeliscidae 

Ampelisca anomala Sars 86 10 

Ampelisca brevicornis (Costa) 69 1 

Ampelisca byblisoides K. H. Barnard 53 2 

103 1 

Ampelisca palmata K. H. Barnard 53 3 

Byblis anisuropus Stebbing 86 1 

Byblis gaimardi (Kroyer) 60 1 

109 3 

Family Amphilochidae 

Hoplopleon medusarum K. H. Barnard 103 1 

Unguja yaya Griffiths 86 1 

Family Corophiidae 

Chevalia aviculae Walker 86 77 

103 2 

Concholestes armatus Griffiths . 31 3 

78 31 

86 4 

103 34 

109 13 

Gammaropsis afra Stebbing 86 41 

91 3 

103 32 

109 2 

Photis uncinata K. H. Barnard 86 26 

Unciolella spinosa Griffiths 60 1 

69 1 

86 8 

109 1 

Family Dexaminidae 

Atylus homochir Haswell 109 1 

Lepechinella occlo J. L. Barnard 60 1 

Family Eusiridae 86 9 

Dautzenbergia grandimanus (Chevreux) 60 1 

107 4 

Family Gammaridae 

Maera inaequipes (Costa) ........ 86 6 

Maera mastersi (Haswell) 86 2 

Family Haustoriidae 

Urothoe elegans Bate 53 1 

Urothoides inops J. L. Barnard 86 1 

Family Ischyroceridae 

Cerapus tubularis Say 103 26 

Family Leucothoidae 

Leucothoe dolichoceras K. H. Barnard 103 2 

Leucothoe spinicarpa (Abildgaard) 86 12 



THE SOUTH AFRICAN MUSEUMS MEIRING NAUDE CRUISES 107 

SM Station Number of 
Number Individuals 

Family Liljeborgiidae 

Liljeborgia epistomata K. H. Barnard 86 2 

Family Lysianassidae 

Amaryllis macrophthalma Haswell 86 24 

103 26 

Bathyamaryllis conocephala (K. H. Barnard) .... 60 2 

61 2 

78 1 

103 1 

109 1 

Hippomedon longimanus (Stebbing) 69 1 

86 2 

109 2 

Hippomedon onconotus (Stebbing) 53 1 

Izinkala fihla gen. et sp. nov 86 4 

103 1 

Lepidepecreum clypeatum Chevreux 60 1 

103 2 

Lepidepecreum twalae Griffiths ....... 60 1 

Lysianassa ceratina 69 1 

86 1 

Valettiella castellana gen. et sp. nov 86 1 

Family Oedicerotidae 

Perioculodes longimanus (Bate & Westwood) .... 86 1 

Family Phoxocephalidae 

Heterophoxus opus Griffiths 60 3 

69 3 

86 8 

Family Podoceridae 

Laetmatophilus purus Stebbing 86 2 

Family Stegocephalidae 

Stegocephaloides australis K. H. Barnard .... 86 1 

Family Stenothoidae 

Parametopa grandimana Griffiths 86 13 

103 11 

Proboloides rotunda (Stebbing) 60 1 

103 6 

Superfamily talitroidea 

Family Phliantidae 

Plioplateia triquetra K. H. Barnard 86 2 

103 43 

Suborder Caprellidea 

Family Aeginellidae 

Eupariambus fallax K. H. Barnard 60 3 

69 1 

103 8 

Pseudoprotella phasma (Montagu) 86 7 

Family Phtisicidae 

Phtisica marina Slaber 86 2 



108 



ANNALS OF THE SOUTH AFRICAN MUSEUM 



SYSTEMATIC DISCUSSION 
Family Acanthonotozomatidae 

Dikwa acrania Griffiths, 1974 

Fig. 1 
Dikwa acrania Griffiths, 1974: 266, fig. 2. 

Material 

SM 86, 2 specimens 

Remarks 

This record is only the second of this interesting species and extends its 
range considerably to the north. Further drawings of certain appendages are 
provided here to supplement the original description. These reveal a number of 
interesting features previously unnoticed. The surface of the mandibular molar 
is strongly triturative and projects a considerable distance from the body of the 
mandible, the apical margin is formed into a sharp chitinized ridge. Article 2 of 
pereiopods 3 and 4 is posteriorly keeled on inner and outer margins, leaving a 




Fig. 1. Dikwa acrania Griffiths, 1974. 

Ovigerous female, 4,5 mm. A. Pereiopod 4 with dactyl enlarged. B. Medial view of pereiopod 5. 

C. Mandible. D. Telson. 



THE SOUTH AFRICAN MUSEUMS M El RING NAUDE CRUISES 109 

deep groove posteriorly, article 4 is acutely produced distally and the dactyl 
bears small chitinized teeth. The fifth pereiopod was erroneously described as 
missing in the original description. It is, in fact, grossly reduced with article 2(?) 
apparently fused to, and partially covered by, an enlarged coxa 7; the remainder 
of the appendage consists of two small articles bearing short anterior spines. 
This remarkable feature is somewhat similar to that found in Tetradeion 
(Stegocephalidae) and should be incorporated into the generic definition of 
Dikwa. 

Distribution 

Endemic, Still Bay to Zululand 200-550 m. 

Family Dexaminidae 

Lepechinella occlo J. L. Barnard, 1973 

Fig. 2 

Lepechinella occlo J. L. Barnard, 1973: 21, figs 8-9. 

Material 

SM 60, 1 specimen 
SM 86, 9 specimens 

Remarks 

The present material differs only marginally from that described by 
Barnard. Thus in South African specimens coxae 1 and 2 are more distinctly 
bifid, the accessory teeth on pereon segments 6 and 7 and pleon segments 1-3 
somewhat more distinct, and the epimeral setae less marked. These differences 
cannot be considered of taxonomic significance, particularly in the light of 
similar size-related differences reported by Barnard (1973). 

Distribution 

New Zealand, 721-860 m; Natal 550-810 m. This record is the first from 
southern Africa. 

Family Eusiridae 

Dautzenbergia grandimanus (Chevreux, 1887) 

Fig. 3 

Parapleustes megachir Walker, 1897: 230, pi. 18 (fig. 4). 
Sympleustes megachir: Stebbing, 1906: 317. Stephensen, 1944: 5, fig. 1. 
Sympleustes grandimanus: Sexton, 1909: 857, pi. 90 (figs 8-32). 
Sympleustes {Dautzenbergia) grandimana: K. H. Barnard, 1937: 158. 
Dautzenbergia grandimanus: J. L. Barnard, 1961: 106. 



Material 

SM 60, 1 specimen 
SM 107, 4 specimens 



110 



ANNALS OF THE SOUTH AFRICAN MUSEUM 




Fig. 2. Lepechinella occlo J. L. Barnard, 1973. 

Male, 6 mm. A. Lateral aspect. B. Accessory flagellum. C. Mandible. D-E. Coxae I, 2. 
F. Third pleonal epimeron. G. Uropod 3. H. Telson. 



THF. SOUTH AFRICAN MUSEUM'S M FIRING NA UDE CRUISES 



111 




Fig. 3. Dautzenbergia grandimanus (Chevreux, 1887). 

Ovigerous female, 8,5 mm. A. Upper lip. B. Lower lip. C. Mandible. D. Maxilla 
E. Maxilliped. F-G. Gnathopods 1,2. H. Uropod 3 and telson. T. Telson. 



1 1 2 ANNALS OF THE SOUTH AFRICAN MUSEUM 

Remarks 

The genus Dautzenbergia has recently been revived by J. L. Barnard (1961) 
to accommodate species of 'Sympleustes' with a partially cleft telson. Of the 
three species in the genus D. grandimanus and D. megacheir appear to be 
indistinguishable. The present material correlates closely with the descriptions 
given by Sexton (1909) and Stephensen (1944). 

Distribution 

North Atlantic, Indian Ocean. This is the first record from southern 
Africa. 

Family Haustoriidae 

Urothoides inops J. L. Barnard, 1967 
Fig. 4 
Urothoides inops J. L. Barnard, 1967: 23, figs 8-10. 
Material 

SM 86, 1 specimen 

Remarks 

There can be no doubt that the present material is synonymous with that 
illustrated by J. L. Barnard (1967). This unusual species can readily be dis- 
tinguished by its broad, flat, down-turned rostrum, which covers the anterior 
end of the body and deflects antenna 2 laterally. Pereiopods 3-5 are strongly 
developed and have article 2 thickened by glandular tissue. The mouthparts, and 
particularly the mandibles, are extremely large relative to the body, which is 
almost as broad as long. 

Distribution 

California, 2 700 m; Natal 550 m. This is the first record of this species 
from the southern African region. 

Family Lysianassidae 

Bathyamaryllis conocephala (K. H. Barnard, 1925) 

Fig. 5 

Amaryllis conocephalus K. H. Barnard, 1925: 324. 
Bathyamaryllis conocephala: K. H. Barnard, 1940: 441. 

Material 

SM 61,2 specimens (including neotype SAM-A 13657) 
SM 78, 1 specimen 
SM 109, 1 specimen 
SM 103, fragment 
SM 60, 2 specimens 



THE SOUTH AFRICAN MUSEUMS ME1RING NAUDE CRUISES 



113 




Fig. 4. Uwthoides inops J. L. Barnard, 1967. 

Female, 2 mm. A. Ventral view of rostrum. B. Mandible. C. Lower lip. D. Maxilliped. 
E. Gnathopod 1. F-H. Pereiopods 3, 4, 5. I. Uropod 3. J. Telson. 



114 



ANNALS OF THE SOUTH AFRICAN MUSEUM 




Female, 5 mm. 
E 



Fig. 5. Bathyamaryllis conocephala (K. H. Barnard, 1925). 
A. Lateral aspect. B. Head and gnathopod 1. C. Mandible. D. Maxilla 1 
Maxilliped. F. Gnathopod 2. G-H. Uropods 2, 3. I. Telson. 



THE SOUTH AFRICAN MUSEUM'S MEIRING NAUDE CRUISES 115 

Remarks 

This species has hitherto been known only through K. H. Barnard's 
somewhat scanty and unfigured description of the holotype female. When an 
attempt was made to obtain this specimen from Barnard's type collection it was 
found to have completely decomposed. A neotype has hence been selected from 
the present material and is shown in Figure 5. The bevelled off antero-lateral 
angles of the head clearly identify this species and distinguish it from the allied 
B. rostrata (Chevreux), in which the lateral lobes of the head are large and 
vertically truncated. The unusual form of the head in these two species is quite 
distinct from that in the other members of the genus— B. haswelli (Stebbing) and 
B. perezii Pirlot. 

Distribution 

Endemic, Cape Peninsula to Natal, 680-1 300 m. 

Izinkaia gen. no v. 
Diagnosis 

Head and coxa 1 both very small, largely concealed by enlarged coxa 2; 
epistome and upper lip inconspicuous; mandible with distinct non-dentate 
incisor, palp attached distally; maxillae poorly developed; maxilliped lacking 
plates, palp with minute article 4; gnathopod 1 slender, minutely subchelate, 
articles 3 and 5 elongate; gnathopod 2 chelate; uropod 3 reduced, enclosed 
within urosomite 3, outer ramus with small article 2; telson entire. 

Type species 

Izinkaia fihla sp. nov. 

Relationships 

This remarkable genus appears quite discrete from any known form. The 
greatly reduced mouthparts are of particular note. A comparable degree of 
reduction occurs in Kerguelenia Stebbing, but here the mandible lacks both 
incisor and molar, while the maxilliped retains small plates and a normal palp 
of four articles. The two genera, however, differ grossly in other respects, such 
as the form of gnathopod 1 and relative prominence of coxa 1, although other 
notable similarities with certain species of Kerguelenia occur, for example the 
enclosure of uropod 3 within urosomite 6 in K. borealis Sars. Izinkaia gen. nov. 
also bears a certain resemblance to various genera in the Lepidepeereum group. 
Thus species such as L. clypodentatum J. L. Barnard show carinate peduncular 
segments in antenna 1, similar gnathopods 1 and 2 and enlarged article 2 of 
pereiopod 5. Clepidecrella J. L. Barnard lacks certain of these features but has 
reduced maxillipedal plates, a short uropod 3 and entire telson. Despite these 
features, Izinkaia gen. nov. remains quite distinct with its grossly reduced coxa 1 
and forward projecting, enlarged coxa 2. 



116 ANNALS OF THE SOUTH AFRICAN MUSEUM 

Izinkala fihla sp. no v. 

Figs 6-7 
Description (of ?female 3,2 mm) 

Head shorter than pereon segment 1 , post-antennal angles acutely produced ; 
peduncle of antenna 1 large, article 1 acutely produced, article 2 overhanging 
remainder of antenna as a broad lobe with lateral flanges between which distal 
articles originate, flagellum 4-articulate, article 1 broad, posteriorly fringed with 
long aesthetascs, accessory flagellum 3-articulate; antenna 2 with article 1 sub- 
circular, apparently fused to head, gland cone prominent, flagellum 3-articulate ; 
epistome and upper lip inconspicuous; mandible with non-dentate incisor, spine 
row absent, molar apparently large and smooth; maxillae not satisfactorily 
resolved, minute; maxilliped completely lacking both inner and outer plates, 
palp article 4 reduced. 

Coxa 1 small and thickened by muscle fibres, completely concealed by large 
forward-projecting coxa 2, margins of coxae 2-4 of the two sides abutting 
ventrally, the small gap below the head closed by projecting lobe of article 2 of 
antenna 1 ; gnathopod 1 slender, elongate, articles 3 and 5 each as long as 2, 
palm slightly oblique, defined by a single strong spine which projects into a 
pocket formed within the thick, highly chitinized dactyl, which bears sharp 
transparent lateral flanges; gnathopod 2 chelate; article 2 of pereiopod 3 
rotund, article 4 greatly widened posteriorly; pereiopod 4 similar in structure to 
3 but longer; pereiopod 5 with article 2 elongate and produced distally as far as 
tip of article 4, articles 4 and 5 acutely lobed posteriorly. 

Pleonal epimera 1 and 2 smoothly rounded, 3 rounded-quadrate; pleon 
segment 4 with a rounded dorsal carina overhanging pleon segment 5; uropods 1 
and 2 strongly spinose dorsally, rami equal; pleon segment 6 overhanging 
uropod 3 and telson above and to the sides; uropod 3 very small, outer ramus 
with a spinose article 2 overhung by dorsal projection of article 1, inner ramus 
shorter, evenly tapering; telson elongate, entire, bearing strong dorsal and 
terminal spines. 

Holotype 

SAM-A13659, Tfemale, 3,2 mm. 

Type locality 

SM 86, 27°59,5'S 32°40,8'E, 550 m, 22 May 1976. 

Material 

SM 86, 4 specimens 
SM 103, 1 specimen 

Valettiella gen. no v. 
Diagnosis 

Antenna 1 geniculate, articles 2 and 3 slender and elongate; article 1 of 
antenna 2 greatly enlarged; upper lip and epistome inconspicuous; mandibular 



THE SOUTH AFRICAN MUSEUM'S ME1RING NAUDE CRUISES 



117 




Fig. 6. Izinkala fihla gen. et sp. nov. 

Female 3,2 mm. A. Lateral aspect. B. Antennae 1 and 2. C. Mandible. D. Maxilliped 

E-F. Unidentified mouthparts, probably maxilla 1 . 



118 



ANNALS OF THE SOUTH AFRICAN MUSEUM 




Fig. 7. Izinkala fihla gen. et sp: nov. 

Female, 3,2 mm. A. Gnathopod 1 with palm enlarged. B. Gnathopod 2. C-D. Pereiopods 3, 5. 

E. Lateral view of urosome. F. Uropod 3. G. Telson. 



THE SOUTH AFRICAN MUSEUM'S MH1R1NO NAUDE CRUISES 1 19 

molar triturative, incisor strongly toothed, palp attached level with molar; 
inner plate of maxilla 1 setose medially; coxae 1 and 2 rectangular and equally 
elongate; gnathopod 1 weakly chelate; gnathopod 2 subchelate; telson short, 
cleft. 

Type species 

Valettiella castellana sp. nov. 

Relationships 

The strongly dentate mandibular incisor and unspecialized gnathopods of 
this genus place it in a distinct group presently comprising Valettia Stebbing, 
1888 and Valettiopsis Holmes, 1908. 

The new form, however, shows features intermediate between those 
defining Valettia and Valettiopsis, as well as a number of distinct characteristics 
of its own. Thus Valettia has the outer plate of the maxilliped apically produced 
and coxae 1 and 2 similarly short and broad, while Valettiopsis has a normal 
maxilliped but coxa 2 elongate and almost covering a reduced coxa 1 . Valettiella 
gen. nov. not only combines a normal maxilliped with elongate and equal 
coxa 1 and 2, but also differs from both the above genera in its unusual geniculate 
antenna 1 with elongate peduncle, enlarged article 1 of antenna 2, widened and 
castelloserrate article 2 of posterior pereiopods and shortened uropod 3 with 
reduced inner ramus. 

Valettiella castellana sp. nov. 
Figs 8-9 
Description (of ovigerous female, 4 mm) 

Head as long as first two pereon segments, ocular lobes acute, eyes weak; 
antenna 1 with article 1 elongate, article 2 almost as long as 1 , normally folded 
back into ventral groove of article 1 , flagellum 9-articulate, accessory flagellum 
uni-articulate ; article 1 of antenna 2 acutely produced into an enlarged shield 
projecting just below ocular lobe of head, gland cone prominent, flagellum 
5-articulate ; mandible with 3-articulate palp, incisor with 6 strong teeth, lacinia 
mobilis with 4 teeth, spine row of 8 spines, molar large and strongly ridged; 
maxilla 1 with bi-articulate palp projecting beyond tip of outer plate, palp with 
7 terminal spine teeth, outer plate bearing 8 strong serrate spines, inner plate 
setose medially; inner plate of maxilla 2 strongly setose medially; maxilliped 
with powerful 4-articulate palp, outer plate apically rounded, bearing eight 
medial spine teeth. 

Coxae 1-3 equally elongate, 1 and 2 with small postero-distal tooth; 
gnathopod 1 weakly chelate, article 5 considerably larger than 6; gnathopod 2 
with article 3 only slightly elongate, articles 5 and 6 subequal, palm oblique; 
pereiopods 3-5 progressively longer, article 2 greatly lobed posteriorly, anterior 
margin spinose, posterior margin strongly castelloserrate. 



120 



ANNALS OF THE SOUTH AFRICAN MUSEUM 




Fig. 8. Valettiella castellana gen. et sp. nov. 

Ovigerous female, 4 mm. A. Lateral aspect. B. Antenna 1 with accessory flagellum enlarged. 
C. Antenna 2. D. Mandible. E. Lower lip. F. Maxilla 1. 



THE SOUTH AFRICAN MUSEUM'S MEIRING NAUDE CRUISES 



121 




Fig. 9. Valet tiella castellana gen. et sp. nov. 

Ovigerous female, 4 mm. A. Maxilla 2. B. Maxilliped. C. Gnathopod 1. D. Gnathopod 2 

with palm enlarged. E. Pereiopod 5. F-H. Uropods 1, 2, 3. I. Telson. 



122 



ANNALS OF THE SOUTH AFRICAN MUSEUM 



Pleon segments 1, 2, 4 and 5 each bearing a single medio-dorsal tooth, 
pleonal epimera quadrate postero-distally; rami of uropods 1 and 2 equal, 
strongly spinose dorsally and apically; uropod 3 considerably shorter than 1 
and 2, outer ramus with a small article 2, inner ramus about half size of outer 
and closely appressed to its medio-ventral surface; telson short, cleft, each lobe 
bearing a large terminal spine and three small setae. 

Holotype 

SAM-A 13658, ovigerous female, 4 mm, unique. 

Type locality 

SM 86, 27°59,5'S 32°40,8'E, 550 m, 22 May 1976. 

Family Stenothoidae 

Parametopa grandimana Griffiths, 1974 
Fig. 10 
Parametopa grandimana Griffiths, 1974: 324, fig. 18. 

Material 

SM 86, 13 specimens 
SM 103, 1 1 specimens 




Fig. 10. Parametopa grandimana Griffiths, 1974. 
Male, 4,5 mm. A. Gnathopod 1. B. Gnathopod 2. 



THE SOUTH AFRICAN MUSEUM'S MEIR1NG NAUDE CRUISES 123 

Remarks 

The above material includes adult males of up to 4,5 mm which show 
gnathopods somewhat different to those described by Griffiths (1974). In 
terminal males the palm of gnathopod 1 is defined by two small spines and the 
dactyl bears three setae. Article 6 of gnathopod 2 is somewhat more elongate 
than is typical of smaller males and the palm is undefined, while the dactyl 
becomes greatly thickened terminally. 

Distribution 

Endemic, Still Bay to Natal 200-680 m. 

ACKNOWLEDGEMENTS 

This project was made possible through the generosity of the staff of the 
South African Museum, who not only lent the author the material but also 
provided excellent library facilities. I am indebted to the South African Council 
for Scientific and Industrial Research for financial support. My thanks also to 
Mrs S. Hardman who kindly typed the manuscript. 

REFERENCES 

Barnard, J. L. 1961. Gammaridean Amphipoda from depths of 400 to 6000 meters. 

Galathea Rep. 5: 23-128. 
Barnard, J. L. 1967. Bathyal and abyssal gammaridean Amphipoda of Cedros Trench, Baja 

California. Bull. U.S. natn. Mus. 260: 1-205. 
Barnard, J. L. 1973. Deep-sea Amphipoda of the genus Lepechinella (Crustacea). Smithson. 

Contr.Zool. 133: 1-31. 
Barnard, K. H. 1925. Contributions to the crustacean fauna of South Africa. 8. Further 

additions to the list of Amphipoda. Ann. S. Afr. Mus. 20: 319-380. 
Barnard, K. H. 1937. Amphipoda. Scient. Rep. John Murray Exped. 1933-34 5: 131-201. 
Barnard, K. H. 1940. Contributions to the crustacean fauna of South Africa. 12. Further 

additions to the Tanaidacea, Isopoda, and Amphipoda with keys for the identification of 

hitherto recorded marine and freshwater species. Ann. S. Afr. Mus. 32: 381-543. 
Griffiths, C. L. 1974. The Amphipoda of southern Africa, Part 4. The Gammaridea and 

Caprellidea of the Cape Province east of Cape Agulhas. Ann. S. Afr. Mus. 64: 251-336. 
Louw, E. 1977. The South African Museum's Cruises on the R/V Meiring Naude off the east 

coast of southern Africa. 1. Station data, 1975, 76. Ann. S. Afr. Mus. 8: 147-159. 
Stebbing, T. R. R. 1906. Amphipoda 1. Gammaridea. Tierreich 21: I-XXXIX, 1-806. 
Sexton, E. W. 1909. Notes on some Amphipoda from the north side of the Bay of Biscay: 

Families Pleustidae and Eusiridae. Proc. zool. Soc. Lond. 1909: 848-879. 
Stephensen, K. 1944. Crustacea Malacostraca, VIII: (Amphipoda IV). Danish Ingolf-Exped. 

3: 1-51. 
Walker, A. O. 1897. On some new species of Edriophthalma from the Irish Seas. J. Linn. Soc. 

26: 226-232. 



6. SYSTEMATIC papers must conform to the International code of zoological nomenclature 
(particularly Articles 22 and 51). 

Names of new taxa, combinations, synonyms, etc., when used for the first time, must be 
followed by the appropriate Latin (not English) abbreviation, e.g. gen. nov., sp. nov., comb, 
nov., syn. nov., etc. 

An author's name when cited must follow the name of the taxon without intervening 
punctuation and not be abbreviated; if the year is added, a comma must separate author's 
name and year. The author's name (and date, if cited) must be placed in parentheses if a 
species or subspecies is transferred from its original genus. The name of a subsequent user of 
a scientific name must be separated from the scientific name by a colon. 

Synonymy arrangement should be according to chronology of names, i.e. all published 
scientific names by which the species previously has been designated are listed in chronological 
order, with all references to that name following in chronological order, e.g. : 

Family Nuculanidae 

Nuculana {Lembulus) bicuspidata (Gould, 1845 

Figs 14-15A 
Nucula (Leda) bicuspidata Gould, 1845: 37. 
Leda plicifera A. Adams, 1856: 50. 

Laeda bicuspidata Hanley, 1859: 118, pi. 228 (fig. 73). Sowerby, 1871 : pi. 2 (fig. 8a-b). 
Nucula largillierti Philippi, 1861: 87. 
Leda bicuspidata: Nickles, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9. 

Note punctuation in the above example : 

comma separates author's name and year 

semicolon separates more than one reference by the same author 

full stop separates references by different authors 

figures of plates are enclosed in parentheses to distinguish them from text-figures 

dash, not comma, separates consecutive numbers 

Synonymy arrangement according to chronology of bibliographic references, whereby 
the year is placed in front of each entry, and the synonym repeated in full for each entry, is 
not acceptable. 

In describing new species, one specimen must be designated as the holotype; other speci- 
mens mentioned in the original description are to be designated paratypes ; additional material 
not regarded as paratypes should be listed separately. The complete data (registration number, 
depository, description of specimen, locality, collector, date) of the holotype and paratypes 
must be recorded, e.g. : 

Holotype 

SAM-A13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King's Beach 
Port Elizabeth (33°51'S 25°39'E), collected by A. Smith, 15 January 1973. 

Note standard form of writing South African Museum registration numbers and date. 

7. SPECIAL HOUSE RULES 

Capital initial letters 

(a) The Figures, Maps and Tables of the paper when referred to n the text 

e.g. '. . . the Figure depicting C. namacolus . . .'; '. . . in C. namacolus (Fig. 10) . . .' 

(b) The prefixes of prefixed surnames in all languages, when used in the text, if not preceded 
by initials or full names 

e.g. Du Toit but A. L. duToit; Von Huene but F. von Huene 

(c) Scientific names, but not their vernacular derivatives 
e.g. Therocephalia, but therocephalian 

Punctuation should be loose, omitting all not strictly necessary 
Reference to the author should be expressed in the third person 
Roman numerals should be converted to arabic, except when forming part of the title of a 

book or article, such as 

'Revision of the Crustacea. Part VIII. The Amphipoda.' 
Specific name must not stand alone, but be preceded by the generic name or its abbreviation 

to initial capital letter, provided the same generic name is used consecutively. 
Name of new genus or species is not to be included in the title : it should be included in the 

abstract, counter to Recommendation 23 of the Code, to meet the requirements of 

Biological Abstracts. 



CHARLES GRIFFITHS 

THE SOUTH AFRICAN MUSEUM'S 
MEIRING NAUDE CRUISES 

PART 6 
AMPHIPODA 



VOLUME 74 PART 5 



JANUARY 1978 



ISSN 0303-2515 



MUS. COMP. ZOOL. 
LIBRARY 

APR 2 8 

HARVAF 
UN! V 



ANNALS 




GAPE TOWN 



INSTRUCTIONS TO AUTHORS 

1. MATERIAL should be original and not published elsewhere, in whole or in part. 

2. LAYOUT should be as follows: 

(a) Centred masthead to consist of 

Title: informative but concise, without abbreviations and not including the names of new genera or species 

Author's(s') name(s) 

Address(es) of author(s) (institution where work was carried out) 

Number of illustrations (figures, enumerated maps and tables, in this order) 

(b) Abstract of not more than 200 words, intelligible to the reader without reference to the text 

(c) Table of contents giving hierarchy of headings and subheadings 

(d) Introduction 

(e) Subject-matter of the paper, divided into sections to correspond with those given in table of contents 

(f) Summary, if paper is lengthy 

(g) Acknowledgements 
(h) References 

(i) Abbreviations, where these are numerous 

3. MANUSCRIPT, to be submitted in triplicate, should be typewritten and neat, double spaced 
with 2,5 cm margins all round. First lines of paragraphs should be indented. Tables and a list of 
legends for illustrations should be typed separately, their positions indicated in the text. All 
pages should be numbered consecutively. 

Major headings of the paper are centred capitals; first subheadings are shouldered small 
capitals; second subheadings are shouldered italics; third subheadings are indented, shouldered 
italics. Further subdivisions should be avoided, as also enumeration (never roman numerals) 
of headings and abbreviations. 

Footnotes should be avoided unless they are short and essential. 

Only generic and specific names should be underlined to indicate italics; all other marking 
up should be left to editor and publisher. 

4. ILLUSTRATIONS should be reducible to a size not exceeding 12 x 18 cm (19 cm including 
legend); the reduction or enlargement required should be indicated; originals larger than 
35 x 47 cm should not be submitted; photographs should be rectangular in shape and final 
size. A metric scale should appear with all illustrations, otherwise magnification or reduction 
should be given in the legend ; if the latter, then the final reduction or enlargement should be 
taken into consideration. 

All illustrations, whether line drawings or photographs, should be termed figures (plates 
are not printed; half-tones will appear in their proper place in the text) and numbered in a 
single series. Items of composite figures should be designated by capital letters; lettering of 
figures is not set in type and should be in lower-case letters. 

The number of the figure should be lightly marked in pencil on the back of each illustration. 

5. REFERENCES cited in text and synonymies should all be included in the list at the end of 
the paper, using the Harvard System {ibid., idem, he. cit., op. cit. are not acceptable): 

(a) Author's name and year of publication given in text, e.g. : 

'Smith (1969) describes . . .' 

'Smith (1969: 36, fig. 16) describes ..." 

'As described (Smith 1969a, 19696; Jones 1971)' 

'As described (Haughton & Broom 1927) . . .' 

'As described (Haughton et al. 1927) . . .' 

Note: no comma separating name and year 

pagination indicated by colon, not p. 

names of joint authors connected by ampersand 

et al. in text for more than two joint authors, but names of all authors given in list of references. 

(b) Full references at the end of the paper, arranged alphabetically by names, chronologically 
within each name, with suffixes a, b, etc. to the year for more than one paper by the same 
author in that year, e.g. Smith (1969a, 19696) and not Smith (1969, 1969a). 

For books give title in italics, edition, volume number, place of publication, publisher. 

For journal article give title of article, title of journal in italics (abbreviated according to the World list o y 
scientific periodicals. 4th ed. London: Butterworths, 1963), series in parentheses, volume number, part 
number (only if independently paged) in parentheses, pagination (first and last pages of article). 

Examples (note capitalization and punctuation) 

Bullough, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan. 

Fischer, P.-H. 1948. Donnees sur la resistance et de le vitalite des mollusques. /. Conch., Paris 88: 100-140. 

Fischer, P.-H., Duval, M. & Raffy, A. 1933. Etudes sur les echanges respiratoires des littorines. Archs 

Zool. exp. gen. 74: 627-634. 
Kohn, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon. 

Ann. Mag. nat. Hist. (13) 2: 309-320. 
Kohn, A. J. 19606. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean. 

Bull. Bingham oceanogr. Coll. 17 (4): 1-51. 
Thiele, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. In: Schultze, L. Zoologische 

und anthropologische Ergebnisse einer Forschungsreise im west lichen und zentralen Sud-Afrika 4: 269-270. 

Jena: Fischer. Denkschr. med.-naturw. Ges. Jena 16: 269-270. 

(continued inside back cover) 



ANNALS OF THE SOUTH AFRICAN MUSEUM 
ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM 



Volume 74 Band 
January 1978 Januarie 
Part 5 Deel 




THE SOUTH AFRICAN MUSEUM'S 

MEIRING NAUDE CRUISES 

PART 7 

MARINE ISOPODA 



By 



BRIAN KENSLEY 



Cape Town Kaapstad 



The ANNALS OF THE SOUTH AFRICAN MUSEUM 

are issued in parts at irregular intervals as material 
becomes available 

Obtainable from the South African Museum, P.O. Box 61, Cape Town 



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Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap 



THE SOUTH AFRICAN MUSEUM'S M EI RING NAUDE CRUISES 

PART 7 
MARINE ISOPODA 

By 

Brian Kensley 

South African Museum, Cape Town 

(With 21 figures and 1 table) 

[MS. accepted 9 August 1977] 

ABSTRACT 

Excluding the Anthuridea, twenty-one species of isopods are recorded from deep water 
off the Natal coast, including fourteen new records. The latter include one new genus, 
Spinarcturus, and ten new species. These are Microarcturus biserialis, M. youngi, Neastacilla 
longispina, Spinarcturus natalensis, Excirolana bicornis, Cirolana caeca, Serolis brinki, 
Haploniscus gernekei, Eurycope glabra, and Ilyarachna wolffi. Five endemic South African 
species are also recorded, as well as Bathycopea typhlops, previously only known from the 
coast of Ireland. 



CONTENTS 



Introduction 
Species list . 
Systematic discussion 
General discussion 
Acknowledgements . 
References . 



PAGE 

125 
127 
128 
155 
157 
157 



INTRODUCTION 

The present paper forms part of a series, based on material collected by 
the South African Museum during the 1975, 1976 cruises off the Natal coast on 
the R/V Meiring Naude. For the scope of, and the background to this pro- 
gramme, as well as for all station data, the reader is referred to Louw (1977). 

The Anthuridea, of which about seven species were collected, are not dealt 
with, and will form the subject of a future paper. 

The following abbreviations are used throughout this paper: SAM— South 
African Museum catalogue number; SM— Meiring Naude Station numbers; 
TL— total length; ovig.— ovigerous; juv.— juvenile. 

125 
Ann. S. Afr. Mus. 74 (5), 1978: 125-157, 21 figs, 1 table. 



126 



ANNALS OF THE SOUTH AFRICAN MUSEUM 



31° 10 2ff 30' 40' 50' 32° 10' 2ff 3ff 40' 50' 33° Iff 2ff 




50' 31° 10' 20 30' 40' 5ff 32 s 10' . 20' 30' 40' 50' 330 Iff 20 



Fig. 1. Map showing Meiring Naude 1975, 1976 stations. 



THE SOUTH AFRICAN MUSEUM'S MEIRING NAUDE CRUISES 



127 



SPECIES LIST 

Material not identified to specific level is either damaged or immature. 



SUBORDER VALVIFERA 

Family Arcturidae 
*Microarcturus biserialis sp. nov. 
Microarcturus oudops (Barnard) . 
*Microarcturus youngi sp. nov. . 

*Neastacilla longispina sp. nov. . 
*Spinarcturus natalensis gen. et sp. nov. 

SUBORDER FLABELLIFERA 

Family Cirolanidae 

*Excirolana bicornis sp. nov. 

*Cirolana caeca sp. nov 

Cirolana imposita Barnard .... 

Family Sphaeromatidae 

*Bathycopea typhlops Tattersall . 

Family Serolidae 

*Serolis brinki sp. nov. . . 

Family Cymothoidae 

Nerocila sp 107 

SUBORDER GNATHIIDEA 

Family Gnathiidae 

Gnathia sp .86 

SUBORDER ASELLOTA 

Family Stenetriidae 

^Stenetrium abyssale Wolff 86 

Stenetrium dalmeida Barnard . . . . 86 
Stenetrium dagama Barnard 

Family Haploniscidae 

*Haploniscus gemekei sp. nov 86 

103 

Family Ischnomesidae 

Ischnomesis sp 6 

Family Eurycopidae 

*Eurycope glabra sp. nov 60 

103 

Family Dendrotionidae 

* Acanthomunna spinipes (Vanhoffen) ... 86 

103 

Family Ilyarachnidae 

*Ilyarachna wolffi sp. nov 86 

103 
Ilyarachna sp. 103 

*New Record 



SM 










Station 


3S 


9$ 


ovig. ?? 


Juvs 


109 


3 








103 


2 


4 


4 


— 


86 


12 


5 


3 


±50 


103 


8 


10 


— 


37 


86 


2 


— 


3 


3 


86 


4 


2 


3 


2 


103 


1 


— 


— 


2 


86 


4 


6 




7 


78 


2 


4 


4 


— 


16 


1 


— 


— 


— 


23 


1 


— 


— 


— 


103 


2 


3 


1 


— 


103 


2 


3 


1 





±20 ±20 ±20 ±20 



1 — — 



2 fragments 

- 1 

1 — 

4 8 

2 1 

4 — 

2 — 



2 — — 



128 ANNALS OF THE SOUTH AFRICAN MUSEUM 

SYSTEMATIC DISCUSSION 
Family Arcturidae 

Microarcturus biserialis sp. no v. 

Fig. 2 
Description 

Body elongate, slender, widest at pereionites II and III; head fused with 
pereionite I, anterior margin concave, anterolateral corners acute, eyes lacking, 
one pair small submedian tubercles anterior to submedian pair dorsal spinose 
processes; lateral margin of head incised. Coxae of pereionites II and III acute, 
of pereionites III to VII with two or three smaller points. Each pereionite 
possessing dorsal submedian pair of spinose processes and broader apically 
acute lateral process. Pleon consisting of three fused segments plus pleotelson, 
former each marked by a pair of submedian spines ; pleotelson apically acute, 
with acute lateral tooth at about midpoint. 

Antennule not reaching midpoint of second antennal peduncle segment. 

Three distal segments of antennal peduncle becoming progressively longer. 

Mandibles, maxillae and maxilliped typical of the genus. 

Pereiopod I shorter than following pereiopods, with numerous serrate 
spine-like setae. 

Pereiopods II to IV slender, all segments with elongate simple setae on 
ventral surface. 

Pereiopods V to VII more robust, dactyli, propodi, carpi, meri and distal 
part of ischia with dense ventral pile of short setules, propodi and carpi with few 
fringed ventral spines. 

Penial process with distal third of rami separate, tips truncate. 

Pleopod 1 with endopod about half length of exopod, distally rounded, 
bearing simple setules; exopod apically curved, with only two distal fringed 
setae; about twenty simple spines on outer margin, broad furrow on anterior 
face. Pleopod 2 rami similar, distally rounded/truncate, stylet on endopod only 
slightly longer than ramus, proximally stout, tapering distally, with small spines 
on inturned distal flanges. 

Material 

Holotype SAM-A 15467 1 ^ TL 5,9 mm SM 109 28°41,0'S 32°36,8'E 

1 300 m 
2 SS TL 5,9 mm SM 109 

Remarks 

The structure of the male pleopods 1 and 2, the mouthparts, and pereional 
appendages leave no doubt that the present species belongs to the genus 
Microarcturus, in spite of the lack of eyes and the relatively short exopod of 
pleopod 1. The dorsal sculpturing resembles that found in the males of 
M. ornatus Kensley, 1975, and M. similis (Barnard), but neither of these has 



THE SOUTH AFRICAN MUSEUM'S ME1RING NA UDE CRUISES 



129 




Fig. 2. Microarcturus biserialis sp. nov. 

A. <J dorsal view. B. $ lateral view. C. Pleopod 1 <$. D. Maxilliped. E. Pleopod 2 <$. 
F. Pereiopod I. G. Pereipod II. H. Pereiopod VII. I. Penis. 



130 



ANNALS OF THE SOUTH AFRICAN MUSEUM 



the double row of spines as well developed on the last three pereionites. 

The specific name 'biserialis' is derived from the double row of prominent 
dorsal spines on the pereion and pleon. 

Microarcturus oudops (Barnard) 
Fig. 3 

Neoarcturus oudops Barnard, 1914: 214, pis 18c, 19b; 1920: 397; 1940: 508. Nordenstam, 
1933: 115. 

Previous records 
Off Cape Point. 

Material 

SAM-A15472 2 &} 4 $? 4 ovig. ?? SM 103 

Remarks 

Barnard (1914) based the new genus Neoarcturus on several features, 
including the non-geniculate body, the non-elongate fourth pereionite, distinct 
coxae, pleon of four fused segments, and the antennal flagellum of three articles. 
All these features apply equally to Microarcturus. Kensley (1975) characterized 




Fig. 3. Microarcturus oudops (Barnard) Pleopod 1 &. 



THE SOUTH AFRICAN MUSEUM'S MEIR1NG NAUDE CRUISES 131 

Microarcturus by incised lateral margins of the head, dorsolateral eyes, first 
pereionite completely fused to head, two or three pleonal segments anterior to 
the pleotelson, and the first pleopod of the male with the inner ramus more than 
half the length of the outer. All these features are found in M. oudops. Further, 
although Barnard characterized Neoarcturus as being blind and possessing three 
pairs of oostegites, four pairs of oostegites are present, as are eyes ; these latter 
are, however, very weakly pigmented. There is thus no justification for placing 
oudops in a genus separate from Microarcturus. The relative length of the inner 
ramus of the first pleopod of the male cannot be used as a criterion for generic 
separation as this shows a range from short in M. biserialis to fairly long in 
M. youngi. The first pleopod of this species is refigured. 

Microarcturus youngi sp. no v. 
Fig. 4 

Description 

Female. Body with numerous small tubercles; widest at pereionites II 
and III. Head with anterolateral corners subacute, with submedian ridges. 
Eyes large, lateral. Coxae indistinct, of pereionite I ventrally directed, tridentate; 
coxae of pereionites II and III expanded laterally, convex in outline ; coxa of 
pereionite IV rectangular. Pereionites V to VII shorter and narrower than 
preceding ones. Pleon consisting of one indistinct and two distinct segments 
plus pleotelson. Latter broad, terminally subacute, with rounded lateral tooth. 

Antennule reaching to midpoint of second antenna! peduncle segment. 
Antennal peduncle 4-segmented, two basal segments together equal in length to 
third segment, latter slender, slightly shorter than fourth segment; flagellum of 
two articles with slender terminal spine. 

Mandibles, maxillae and maxilliped typical of the genus. 

Pereiopod I shorter than following pereiopods, with numerous serrate 
spine-like setae. 

Pereiopods II to IV relatively slender, with numerous slender setae. 

Pereiopods V to VII more robust, propodi, carpi and meri with dense very 
short setae on ventral surface plus several widely spaced spines, those on 
propodus barbed. 

Male. Body parallel-sided, cylindrical, pereionites varying in sculpture 
from granular to almost smooth. Pleon with pair of large subcorneal processes 
at base. 

Penial rami fused for three-quarters of length, tips diverging. Pleopod 1 
basis with rounded proximal flange, and fourteen peg-like processes; inner 
ramus shorter than outer, distally rounded/truncate; outer ramus with apex 
produced and bent, numerous spines on outer margin, broad furrow on anterior 
face, ten plumose setae distally. 

Pleopod 2 with stylet of endopod stout, cylindrical, slightly convoluted 
apically. 



132 



ANNALS OF THE SOUTH AFRICAN MUSEUM 




Fig. 4. Microarcturus youngi sp. nov. 

A. $ dorsal view. B. <J lateral view. C. Penis. D. Maxilliped. E. Pleopod 1 $. F. Antennule. 
G. Pereiopod VII. H. Pereiopod II. I. Pleopod 1 $. 



THE SOUTH AFRICAN MUSEUM'S MEJRING NAUDE CRUISES 



133 



Material 

Holotype SAM-A 15465 

Allotype SAM-A 15465 

SAM-A 15466 



1 ovig. ? TL 5,8 mm SM 86 27°59,5'S 

32°40,8'E 550 m 
1 S TL 6,0 mm SM 86 
12 &? 5 $? 3 ovig. $$ 50 juvs SM 86 
8 $$ 10 $? 37 juvs SM 103 



Remarks 

M. youngi is obviously closely related to the group of species including 
M. ornatus Kensley, M. dayi Kensley, and M. similis (Barnard), in which the 
basic plan and sculpture of the males and females are similar. Each, however, 
may be distinguished by details of the sculpture. The most distinctive feature of 
M . youngi is the very prominent pair of bosses at the base of the pleotelson in 
the male. 

The species is named for Mr D. Young, first officer of the R/V Meiring 
Naude. 

Neastacilla longispina sp. no v. 

Figs 5-6 
Description 

Female. Body slender, elongate. Head and first pereionite fused, fusion 
indicated by shallow furrow; eyes large, lateral; pair of dorsal spines and few 




Fig. 5. Neastacilla longispina sp. nov. 
A. Ovigerous $ lateral view. B. <$ lateral view. 



134 



ANNALS OF THE SOUTH AFRICAN MUSEUM 



small tubercles present. Pereionites I and II with small dorsal spine and spinose 
lateral process. Pereionite II with single very strong dorsal spine, smaller spine 
between dorsal spine and spinose lateral process. Pereionite IV elongate, with 
pair of strong submedian dorsal spines at midpoint, and single slightly hooked 
median dorsal spine on posterior margin. Pereionite V with few small tubercles 
and spinose coxa. Pleon consisting of three fused pleonites plus pleotelson, 
segments indicated by shallow furrows; spinose lateral process on third pleonite. 
Pleotelson with distal half tapering, apically acutely rounded. Antennule 
consisting of 3-segmented peduncle, basal segment with small rounded dorsal 
process; flagellum of single article reaching beyond basal antennal segment. 

Antenna longer than the total body length, basal segment short, with 
spinose distal process; following three segments very elongate, slender, flagellum 




Fig. 6. Neastacilla longispina sp. nov. 

Pereiopod VII. B. Pereiopod I. C. Pereiopod II. D. Pleopod 2 $. E. Antennule $. 

F. Pleotelson <$. 



THE SOUTH AFRICAN MUSEUM'S MEIRING NAUDE CRUISES 135 

short, of three articles, terminal article very short. 

Mouthparts typical of the genus. 

Pereiopod I shorter than following pereiopods, dactylus distally rounded/ 
truncate, with strong terminal serrate spine; all segments bearing numerous 
simple setae; serrate spines also present on dactylus and propodus. 

Pereiopods II to IV slender, elongate, dactyli absent, remaining segments 
bearing elongate simple setae. 

Pereiopods V to VII becoming shorter posteriorly, more robust than 
preceding pereiopods, dactyli with two short apical claws, numerous fine setules 
on all segments except basis, especially concentrated on ventral surfaces. 

Male. Pereionite IV more elongate than in female; head and pereionites 
with few scattered tubercles; no elongate dorsal spines as in female. Antennular 
flagellum relatively longer than in female. 

Antenna with second peduncular segment with small scattered tubercles. 

Pleotelson with slight point at lateral midpoint. 

Pleopod 1 with rami of equal length, with elongate distal plumose setae; 
exopod with three elongate proximal setae, endopod with proximal indentation. 

Pleopod 2 rami of equal length, with distal elongate plumose setae; 
endopod with stylet at base of inner margin extending well beyond rami, 
apically acute and slender. 

Material 

Holotype SAM-A 15459 1 S TL 10,5 mm SM 86 27°59,5'S 32°40,8'E 

550 m 
Allotype SAM-A15459 1? TL 8,3 mm SM 86 

1 3 2 ?? 3 juvs SM 86 

Remarks 

The present species bears little resemblance to the three species of Neastacilla 
hitherto recorded from South African waters, but most closely resembles 
Astacilla attenuata Hale, 1946, from New South Wales. N. longispina differs 
from the Australian species especially in possessing dorsal elongate spines on 
the pereion, and in having more elongate antennae. 

The specific name is derived from the dorsal spines of the female. 

Spinarcturus gen. no v. 
Diagnosis 

Arcturid possessing a geniculate body; first pereionite not fused with the 
head; dorsolateral eyes; antennae shorter than the total body length; fourth 
pereionite not elongate; anterior four pairs of pereiopods unspecialized, not 
differentiated from the posterior three pairs; pleopod 1 of the male relatively 
unspecialized, not furrowed on anterior face ; stylet of pleopod 2 of the male 
apically simple. 

Type species of the genus Spinarcturus natalensis. 

The generic^rame is derived from the numerous spines of the body. 



136 



ANNALS OF THE SOUTH AFRICAN MUSEUM 



Spinarcturus natalensis sp. no v. 

Figs 7-8 
Description 

Male. Body elongate, geniculate, with numerous short fine setules 
entrapping debris. Head with concave anterior margin, not fused with 
pereionite I. Latter bearing pair of slender spines. Pereionites II to IV with two 
pairs of dorsolateral spines. Pleon consisting of three fused segments, first 
pleonite unarmed, second pleonite with one pair dorsolateral spines, third 
pleonite with pair strong proximolateral spinose processes, rest of segment 
smooth, convex, terminally bluntly rounded. Coxae not dorsally visible. Eyes 
dorsolateral. 

Antennule 4-segmented, terminal article elongate, bearing aesthetascs. 

Antenna shorter than body, peduncle consisting of two short proximal 
segments and two more elongate distal segments, plus flagellum of three articles, 
slender terminal spine present; all segments with scattered setae. Mandible with 
tridentate incisor, tridentate lacinia, four setae in row, broad truncate molar 
process with four stout spines on ventral margin. Maxilla 1 with three fringed 
setae on inner ramus, several simple spines on outer ramus. 

Maxilla 2 with both lobes of outer ramus tipped with two sparsely fringed 





Fig. 7. Spinarcturus natalensis gen. et sp. nov. 
A. c? dorsal view. B. Pleopod 1 $. C. Pleopod 2 <J. D. Penis. 



THE SOUTH AFRICAN MUSEUM'S MEIRING NAUDE CRUISES 



137 



setae, inner ramus with about ten fringed setae. 

Maxilliped with several fringed setae on four distal palp segments and on a 
endite. 

Pereiopod I shorter than following pereiopods, dactylus with strong 
terminal spine, smaller secondary spine, and several fringed spines, propodus 
with many fringed spines; carpus, merus, and ischium with few ventral fringed 
spines and numerous fine short setules. 




Fig. 8. Spinarcturus natalensis gen. et sp. nov. 

A. Pereiopod VII. B. Maxilla 2. C. Mandible. D. Uropod apex. E. Pereiopod I. F. Maxilla \ 

G. Maxilliped. H. Antennule. I. Antenna. 



1 38 ANNALS OF THE SOUTH AFRICAN MUSEUM 

Pereiopods II to IV essentially similar to pereiopods V to VII, latter with 
more elongate propodi and carpi. Pereiopods more or less covered with fine 
short setules and adhering debris. 

Rami of penis united almost to apex, latter tapered, finely setose. 

Pleopod 1 relatively unspecialized, with proximal indentation in endopod, 
both rami apically rounded, with numerous plumose setae. 

Pleopod 2 both rami apically rounded, with plumose setae; stylet of 
endopod sabre-shaped, almost twice length of ramus, apically acute. 

Uropod with strong ridge near outer margin, outer ramus small, triangular, 
inner ramus less than half width of outer, with two subapical spines. 

Female. Similar to male, but pereionites II to IV wider; broodpouch 
formed by four pairs of oostegites. 

Material 

Holotype SAM-A 15473 1 $ TL 6,0 mm SM 86 27°59,5'S 32°40,8'E 

550 m 
Allotype SAM-A 15473 1 ovig. ? TL 8,2 mm SM 86 

3 cJcJ 2 ?$ 1 ovig. $ 2 juvs SM 86 
SAM-A15474 1 J 2 juvs SM 103 

Remarks 

Deciding on the generic position of the present species gives rise to some 
difficulty. Pereionite IV is not elongate, which rules out Astaeilla, Neastacilla, 
Arcturella, Areturina, Arcturopsis, Parastacilla and Arcturinoides. The head is 
not fused with the first pereionite, which eliminates Pleuroprion, Antarcturus 
and Idarcturus. The antenna is shorter than the body, and has a flagellum of 
three articles, which rules out Antarcturus, Arcturus, and Dolichiscus. Pleopod 1 
in the male is not specialized or furrowed as in Austroarcturus, Microarcturus, 
Antarcturus and Holidotea. The elongate stylet of the second pleopod of the 
male is reminiscent of Arcturinoides and Areturina but does not have a trifid 
apex as in these latter. The first pleopod with its proximal indentation again 
resembles that of Areturina and Arcturinoides. The pereiopods, however, are not 
differentiated as they are in these genera. The creation of a new genus for this 
material would thus seem to be warranted. 

The specific name is derived from the Province of Natal. 

Family Cirolanidae 

Excivolana bicornis sp. no v. 

Figs 9-10 
Description 

Male. Body width about half length, widest at pereionites II and VI. Head 
with dorsolateral eyes; two elongate slightly curved apically rounded 'horns' 
anterior to eyes, length of 'horns' varying with total length. Frontal lamina 
distally rounded, dorsally visible between antennal bases, proximally narrowed. 



THE SOUTH AFRICAN MUSEUM'S MEIRING NAUDE CRUISES 



139 




Fig. 9. Excirolana bicornis sp. nov. 

A. (J dorsal view. B. Mandible. C. Maxilla 1. D. Maxilla 2. E. Maxilliped. 

F. Head of <$ in lateral view. 



Clypeus acutely pointed in lateral view. Pereionites unsculptured except for 
faint, impressed line. All coxae pointed, those of pereionites IV to VII being 
elongate-acute. Pleotelson triangular, distal margin dentate, apically acute. 

Antennal peduncle 3-segmented, third segment elongate ; flagellum of eight 
to nine articles. 

Antenna longer than antennule, peduncle 4-segmented, terminal segment 
longest, flagellum of fourteen articles. 

Mandible with 3-segmented palp, middle segment nearly three times length 
of basal segment; incisor 4-dentate; setal row of fourteen to sixteen short 



140 



ANNALS OF THE SOUTH AFRICAN MUSEUM 



spine-like setae; molar process with numerous small teeth on upper surface. 

Maxilla 1 with three setae on inner ramus, ten dentate and simple spines on 
apex of outer ramus. 

Maxilla 2 with several plumose setae on inner ramus, inner lobe of outer 
ramus with five setae, outer lobe with three setae. 

Maxillipedal endite narrow, armed with five plumose setae and single 
coupling hook; palp 5-segmented, third segment longest and broadest, all 
segments setose. 

Pereiopod I with sensory spines on ventral margin of propodus, carpus and 
merus, carpus very short. 

Pereiopod YII armed with clusters of serrate spines on propodus, carpus, 
merus and ischium. 




Fig. 10. Excirolana bicomis sp. nov. 

A. Antenna. B. Antennule. C. Pereiopod I. D. Pereiopod VII. E. Pleopod 2 <J. 

F. Apex of telson. G. Uropod. 



THE SOUTH AFRICAN MUSEUMS MEIRING NAUDE CRUISES 141 

Pleopod 2 with stylet attached to base of endopod, extending well beyond 
both rami, apically tridentate. 

Uropod with inner ramus broader than outer, margins serrate, apically 
narrowed, with several setae and few spines on outer margin. 

Female. Similar to male, but lacking 'horns' on head. 

Material 

Holotype SAM-A 15469 1 £ TL (excl. horns) 5,6 mm SM 86 27°59,5'S 

32°40,8'E 550 m 
Allotype SAM-A15469 1 ? (with oostegites) 5,5 mm SM 86 
ParatypesSAM-A15469 3 &? 5 ?? 7 juvs SM 86 

Remarks 

Sexual dimorphism in Excirolana has been recorded only in one other 
species, E. kumari Bowman, 1971, from Malaysia. In this species the male bears 
a pair of triangular horns at the dorsal posterolateral corners of the head, two 
slender lateral horns on pereionites I and II, plus a mediodorsal spine on 
pereionite II. 

The distinctive pair of horns anterior to the eyes on the head of the male 
(from which is derived the specific name), immediately separates E. bicornis 
from E. kumari. 

Cirolana caeca sp. no v. 

Figs 11-12 
Description 

Body width slightly more than half length, strongly convex dorsally, widest 
at pereionites IV and V, coxae well developed, leaf-shaped ; no dorsal sculpture 
other than impressed line on pereionite VII. Pleon consisting of five segments 
plus pleotelson, lateral margin of first pleonite overlapped by coxae of last 
pereionite; lateral margin of fifth pleonite overlapped by fourth; pleotelson 
wider than long, distally rounded, distal margin crenulate. 

Eyes absent. 

Antennule shorter than antenna, third peduncular segment longest, 
flagellum of five articles. 

Antenna with terminal peduncular segment longest, flagellum of eleven 
articles. 

Mandibular palp 3-segmented, middle segment longest, bearing simple and 
plumose spines; incisor of four broad teeth; setal row of ten curved spine-like 
setae; molar process with marginal row of teeth. 

Maxilla 1 inner ramus short, with three apical plumose setae; outer ramus 
with several dentate and simple spines. 

Maxilla 2 inner ramus with five plumose setae, inner lobe of outer ramus 
with four simple setae, outer lobe with three. 



142 



ANNALS OF THE SOUTH AFRICAN MUSEUM 



Maxillipedal endite short, with one coupling hook and four plumose setae; 
palp 5-segmented, middle segment broadest and longest. 

Pereiopod I shorter than following pereiopods, carpus short and triangular, 
ventral margins of propodus, carpus, and merus bearing few sensory spines. 
Pereiopod VII with serrate spines on ventral and distal margins of segments. 

Penial processes digitiform. 

Pleopod 2 male with stylet on inner ramus longer than rami, apically with 
slender spine and subterminal blunt tooth. 

Uropod with both rami distally narrowly rounded, inner ramus broad, 
with sparse plumose setae. 




Fig. 1 1 . Cirolana caeca sp. nov. 
A. <$ dorsal view. B. Maxilliped. C. Maxilla 1. D. Maxilla 2. E. Mandible. 



THE SOUTH AFRICAN MUSEUM'S MEIRING NAUDE CRUISES 



143 



Material 

Holotype SAM-A 15468 1 $ TL 3,9 mm SM 78 27°31,6'S 32°50,0'E 

750 m 
Allotype SAM-A 15468 1 ovig. ? 4,2 mm SM 78 
Paratypes SAM-A 15468 1 <} TL 3,2 mm 5 ?$ TL 2,8 mm-4,0 mm 

3 ovig. $$ TL 3,6 mm-4,0 mm SM 78 

Remarks 

This very distinctive species differs markedly in body shape from other 
blind species of Cirolana, such as C. californiensis Schultz and C. poissoni 
Monod. 

The specific name is derived from the lack of eyes. 




Fig. 12. Cirolana caeca sp. nov. 

A. Antenna. B. Antennule. C. Pleopod 2 $. D. Uropod. E. Apex of telson. 
F. Pereiopod I. G. Pereiopod VII. 



144 ANNALS OF THE SOUTH AFRICAN MUSEUM 

Family Sphaeromatidae 

Bathycopea typhlops Tattersall 

Fig. 13 

Bathycopea typhlops Tattersall, 1905: 12, 65, pi. 3. Loyola e Silva, 1971 : 216, figs 2-3. Schultz, 
1973:275. 

Previous record 

Off County Kerry, Ireland. 
Material 

SAM-A15461 2 S3 3 $$ 1 ovig. ? 3,9-5,3 mm SM 103 
Remarks 

This is only the second record of this remarkable species. Tattersall (1905) 
recorded B. typhlops from five stations off the Irish coast, in depths ranging from 
about 400 to 830 metres, and in all cases the bottom substrate was fine sand. 
The length of the adult male and female from Ireland was 5,0 mm, agreeing well 
with the present material. No differences in proportions or in the appendages 
could be detected between Tattersalfs and Loyola e Silva's figures and 
descriptions, and the present specimens. The huge distance between these two 
records can only be explained by a lack of sampling of the appropriate substrate. 







Fig. 13. Bathycopea typhlops Tattersall, dorsal view. 

Family Serolidae 

Sevolis brinki sp. no v. 

Figs 14-15 
Description 

Male. Body longer than wide, lacking spines and tubercles on dorsum. 
Head with anterolateral angles acute, slightly produced, anterior margin 



THE SOUTH AFRICAN MUSEUM'S ME1RING NAUDE CRUISES 145 




Fig. 14. Serolis brinki sp. nov. 

medially emarginate, eyes entirely absent. Coxal plates marked off by sutures 
on pereionites II to IV, coxal plate of pereionite VII not extending beyond 
pleural plates of second and third pleonites. Pleotelson with very slight rounded 
median ridge, more strongly indicated proximally, becoming obsolete distally; 
small lateral point at uropodal insertion. Antennular peduncle longer than 
flagellum, latter of twenty articles. 

Antennal peduncle about equal in length to entire antennule, flagellum of 
fourteen articles. 

Left mandible with lacinia tridentate, right mandible with incisor somewhat 
expanded, with two strong spines and rounded tubercles; lacinia process distally 
dentate. 

Maxillipedal palp 3-segmented, distal segment well developed. 

Pereiopod I, palm of propodus with alternating closely packed slender bifid 
and broad finely setulose setae ; distal margin of carpus with two short setae and 
few fine setules. 

Pereiopod II with double row of eight stubby, bristled setae on palm, distal 
margin with several simple setae. 

Pereiopod VII slender, elongate, with setae on all segments except basis. 

Pleopod 2 with distal portion of endopodal stylet about three times longer 
than proximal portion. 

Uropod with exopod well developed, oval, shorter than endopod. 

Colour pattern (when alive): only anterior half of dorsum pigmented 
red-grey; posterior half white. Pigment present on antennal and antennular 
peduncles, with white dapples on head and terga. Head and anterior four 
pereionites with broad hyaline margins. 



146 



ANNALS OF THE SOUTH AFRICAN MUSEUM 



Material 

Holotype SAM-A 15460 1 <? TL 8,6 mm SM 103 28°31,7'S 32°34,0'E 

680 m 
Remarks 

This blind species of Serolis closely resembles two previously recorded 
deep-water species, viz. S. vemae Menzies, 1962, from the North and South 
Atlantic, and S. menziesi Hessler, 1970, from the South Atlantic off Brazil, but 
differs from both. 




Fig. 15. Serolis brinki sp. nov. 

A. Inner view of mandibles. B. Maxilliped. C. Pleopod 2 <$. D. Pereiopod I. 
E. Spines on palm of propodus, pereiopod I. F. Pereiopod II. G. Pereiopod VII. 



THE SOUTH AFRICAN MUSEUM'S ME] RING NAUDE CRUISES 147 

S. menziesi does not have a lateral spine on the pleotelson, the uropodal 
exopod is minute and the pleotelson possesses a triangular flattened proximal 
area. 

In S. vemae the uropodal exopod is also much smaller than in S. brinki, the 
pleotelson also possesses a flattened proximal area, but lacks a lateral spine. 

S. brinki is the first species of the genus recorded in South African waters, 
and is named for Dr C. van der Merwe Brink, President of the South African 
Council for Scientific and Industrial Research. 

Family Stenetriidae 

Stenetrium abyssale Wolff 
Stenetrium abyssale Wolff, 1962: 25, figs 1-5. 

Previous records 

Kermadec Trench (north-east of New Zealand) and Tasman Sea, 4510- 
4540 m. 

Material 

SAM-A15470 1 ovig. $ 5,3 mm 1 # 3,0 mm SM 86 
SAM-A15471 1$ 5,4 mm 4 <?<? 5,0 mm-5,8 mm SM 103 

Remarks 

Wolff (1962) gives the dimensions of 9,9 mm TL $ and 8,6 mm TL $. 
The present specimens, including an ovigerous female, are all less than 6 mm in 
length. This is the only difference from the Kermadec material, the appendages 
and body shape agreeing exactly with Wolff's description. 

Family Haploniscidae 

Haploniscus gernekei sp. no v. 

Fig. 16 
Description 

Body about twice longer than wide. Anterolateral margins of pereionites 
and pleotelson with broad hyaline border. Lateral margin of pleotelson 
continuous with pereion. Distal margin of pleotelson convex, distolateral 
corners not produced beyond medial margin. Rostrum acute, slightly upturned, 
with small tubercle on each side of base ; frontal margin excavate on either side 
of rostrum. Lateral margin of pereionite IV squarely truncate. Pleopod 1 in 
male with rami fused, distal margin straight, with seven setae on each side, 
distolateral portion tapering to narrowly rounded apex. Uropods dorsally 
visible, extending almost to level of medial pleotelsonic margin. 

Material 

Holotype SAM-A 15454 1 <? TL 2,8 mm SM 103 28°31,7'S 32°34,0'E 

680 m 



148 



ANNALS OF THE SOUTH AFRICAN MUSEUM 



Allotype SAM-A 15454 



1 $ SM 103 

8 SS 3 ?$ SM 103 

3^ 3 $$ SM 86 



SAM-A 15455 
Remarks 

H. gemekei differs from H. tricornis Menzies, 



1962, which it closely 
tricornis, five in 
the first pleopod 



resembles, in the antennular flagellum (six articles in H. 
H. gemekei, and the relative lengths of the articles differing) 
of the male, the distolateral angles being more produced than in H. tricornis; 
in the uropod extending at least to the level of the medial pleotelsonic margin 
or slightly beyond, and in lacking a lateral ridge on the dorsum of the pleotelson. 
The species is named for Mr D. Gerneke, previously of the South African 
Museum, for his invaluable assistance on both the Meiring Naude cruises. 




Fig. 16. Haploniscus gemekei sp. nov. 

A. Holotype dorsal view. B. Maxilliped. C. Antennule. D. Apex of pleopod 1 <$. 

E. Pleopod 2 (J. 



THE SOUTH AFRICAN MUSEUM'S MEIRING NAUDE CRUISES 



149 



Family Eurycopidae 

Eurycope glabra sp. nov. 

Figs 17-18 
Description 

Body glabrous, head and first four pereionites narrower than rest of body. 
First four pereionites narrow, anterolateral angles acute ; pereionite V broader 
than VI, lateral corners rounded; pereionite VII wider than VI, distal margin 
slightly concave. Pleotelson of single segment, broader than long, distally 
broadly rounded, uropods just visible dorsally. Rostrum distally rounded, with 
eight setae. 

Basal antennular segment broad, with inner distal angle produced, second 
segment narrow; flagellum of about fifteen articles. 

Mandibular palp with third segment strongly rounded, with several setae 
and fringed spines ; molar process broad, truncate, with five marginal setae. 

Maxilla 1 with several simple curved spines distally, inner ramus with single 
strong distal seta. 

Maxilla 2 slender, with several elongate distal setae. Epipodite of maxilliped 
with strong rounded process on outer margin; three basal palp segments broad, 
second segment longest, with two spines on outer distal angle; two distal 
segments narrow, setose; endite narrower than basal palp segments. 

Pereiopod I very slender, remaining pereiopods missing. 





Fig. 17. Eurycope glabra sp. nov. 
A. Holotype dorsal view. B. Pleopod 1 <$. C. Maxilliped. D. Epipod of maxilliped. 



150 



ANNALS OF THE SOUTH AFRICAN MUSEUM 





Fig. 18. Eurycope glabra sp. nov. 

A. Left mandible. B. Maxilla 1 . C. Maxilla 2. D. Antennule. E. Pleopod 2 <$. F. Uropod. 

G. Rostrum. H. Pereiopod I. 



THE SOUTH AFRICAN MUSEUM'S M El RING NAUDE CRUISES 151 

Pleopod 1 in male narrow, elongate, rami distally rounded, with sub- 
terminal process on outer margin. 

Pleopod 2 with outer margin strongly convex. 

Uropod biramous, inner ramus four and a half times longer than outer. 

Operculum of female with strong median crest, forming spine-like process 
proximally. 

Material 

Holotype SAM-A 15452 1 <J TL 3,6 mm SM 103 28°31,7'S 32°34,0'E 

680 m 

Allotype SAM-A15453 1 ? TL 3,3 mm SM 60 27°09,6'S 32°58,2'E 

800-810 m 
Remarks 

From Wolff's key (1962: 144) to the species of Eurycope the present species 
would seem to resemble E. antarctica VanhofTen, 1914, but this species does not 
have acute antero-lateral corners of the first four pereionites, and the pleotelson 
is not distally as broadly rounded; further, the uropodal exopod is relatively 
longer, the rostrum not as narrow as in the present material. 

The specific name derives from the completely smooth dorsum, entirely 
lacking in sculpture. 

Family Dendrotionidae 

Acanthomunna spinipes (VanhofTen) 
Fig. 19 

Mormomunna spinipes Vanhoffen, 1914: 569, figs 100, 101. 
Acanthomunna spinipes: Menzies, 1962: 174. Wolff, 1962: 65. 

Previous records 

Antarctic (Gauss Station 8.H.1903) 385 m. 

Material 

SAM-A15463 4 &? TL 2,5 mm 8?? 3 ovig. $? TL 4,4 mm SM 86 
SAM-A 15464 2 SS 1? SM 103 

Remarks 

The presence of a uropodal peduncle, pereionites I to III separate, the head 
not fused with pereionite I, maxilliped palp segments all slender, the presence 
of a mobile lacinia in the left mandible, an apically truncate molar process, 
coxae on pereionites II to VII, an antennular flagellum of at least six articles, 
pereiopods with a single claw, and dorsolateral^ inserted uropods, all place 
the present specimens in the family Dendrotionidae. The presence of eyes 
indicate the genus Acanthomunna (Wolff, 1962: 65). The present material agrees 
almost exactly with Vanhoffen's description of A. spinipes, especially in the 
structure of the first and second pleopods of the male. While the generally 
spinose pereion and pleon, and the elongate antennae and pereiopods also 



152 



ANNALS OF THE SOUTH AFRICAN MUSEUM 











Fig. 19. Acanthomunna spinipes (Vanhoffen). 

A. Dorsal view. B. Maxilliped. C. Pleopod 1 $. D. Pleopod 2 $. E. Right mandible. 
F. Maxilla 1. G. Maxilla 2. H. Operculum $. 



THE SOUTH AFRICAN MUSEUM'S ME1R1NG NAUDE CRUISES 153 

agree well, unfortunately the spectacularly spinose uropods of A. spinipes are 
absent in all the Natal material. 

In one specimen, the base of the uropodal peduncle was still in situ. The 
relatively large insertion for the uropods and the sturdy remnant mentioned 
indicate a massive uropod of the type figured by Vanhoffen. 

Family llyarachnidae 

llyarachna wolffi sp. nov. 

Fig. 20 
Description 

Body glabrous, tapering posteriorly. Head with transverse convex portion 
and broad lateral areas. Pereionite I not as broad as II, both laterally acute. 
Pereionites II to IV increasing in length, coxae of pereionites posterior to II 
rounded; posterior margin of pereionite V concave; VI triangular, VII narrower. 
Pleotelson of two segments, terminal segment longer than broad, distally 
narrowly rounded. 

Antennular basal segment broad, outer distal angle somewhat produced, 
second segment narrow; flagellum of about fifteen articles. 

Mandibular palp slender, 3-segmented, middle segment with two fringed 
spines, shorter terminal segment with four spines; incisor process a single 
rounded tooth; molar process narrow, with three terminal setae; strong ridge 
on inner face of mandible. 

Maxilliped with second palp segment very broad and long, third segment 
triangular, two distal segments short and narrow; endite less than half width of 
second palp segment, epipodite broad, distally rounded. 

Pleopod 1 in male curved, rami distally narrowly rounded, setose. 

Uropods missing. 

Material 

Holotype SAM-A 15457 1 # TL 2,9 mm SM 86 27°59,5'S 32°40,8'E 

550 m 
3 33 SM 86 

SAM-A 15458 2 33 SM 103 

Remarks 

Three species closely resemble present material. (See Wolff 1962: 94.) 

/. triangulata Menzies, 1962, is very similar in body proportions but lacks the 
acute coxae of the first two pereionites, the mandibular palp is also very different. 

/. affinis Barnard, 1920, has acute coxae on the first four pereionites, rather 
than just the first two. 

/. crassipes Barnard, 1920, has rounded coxae of the first two pereionites, 
while the pleon is about as long as wide. 

The species is named for Dr Torben Wolff, for his contributions to isopod 
biology and taxonomy. 



154 



ANNALS OF THE SOUTH AFRICAN MUSEUM 




Fig. 20. Ilyarachna woljfi sp. nov. 

A. Holotype dorsal view. B. Pleopod 1 $. C. Maxilliped. D. Antennule. E. Maxilla 1, 
F. Maxilla 2. G. Mandible. H. Pleopod 2 <J. 



THE SOUTH AFRICAN MUSEUM'S MEIRING NA UDE CRUISES 



155 



Uyarachna sp. 

Fig. 21 
Description 

Body markedly spinose. Head with eight spines. Pereionites I to IV with 
two rows of small spines and several lateral spines; pereionites V to VII with 
single row of spines. Pleotelson of two segments, terminal segment with several 
small granules plus single strong pair of spines. 

Material 

SAM-A 15462 2 $$ TL 3,0 mm SM 103 

Remarks 

As the uropoda and all the pereiopods are lacking from both specimens, 
specific status is not given, even though a species of Uyarachna with a double 
row of small spines on the anterior four pereionites does not seem to have been 
recorded. This species is obviously related to such spinose species as 
/. spinosissima Hansen, /. argentinae Menzies, /. gurjanovae Menzies, and 
I. multispinosa Menzies 

GENERAL DISCUSSION 

Of the 21 species of isopod recorded, 15 were taken from either station 
SM 86 or SM 103, 7 species being common to both stations. The nature of the 
bottom at both stations was very similar, and as 9 of the 1 5 species caught are 
new, it may fairly be said that this is a habitat not previously sampled. A 




Fig. 21. Uyarachna sp. ?. 



156 



ANNALS OF THE SOUTH AFRICAN MUSEUM 



comparison of the two stations (Table 1) and the isopods caught at them will 
give some idea of the characteristics of this ecotype. The differences in the catch 
between the stations can to some extent be explained by the difference in fishing 
ability of the gear used. The heavy dredge, with its 100 cm by 30 cm mouth, 
would be able to pick up much larger organisms and rocks than could the light 
biological dredge (see Menzies 1962, fig. 1) with its narrow 100 cm by 10 cm 
mouth. Thus Alcyonaria and Gorgonacea with attendant isopods such as 
Neastacilla would be caught only by the heavy dredge, while some of the smaller 
debris-dwelling isopods of the upper few centimetres of substrate such as 
Eurycope, Serolis and Microarcturus would be caught by the light dredge, but 
perhaps lost through the wider mesh of the heavy dredge. 

Table 1 

A comparison between two stations and the isopods caught at them. 



Depth 

Gear 

Bottom 



Non-Isopod 
organisms 
caught 



Isopods 
caught 



SM 86 

550 metres 
Heavy dredge 

Fine hard mud, overlain with hetero- 
pod and pteropod shells, coral 
fragments, numerous foraminifera, 
sponge fragments and spicules 
Hydroids, gorgonaceans, solitary 
corals, glass sponges, pycnogonids, 
small crabs, cumaceans, amphipods, 
ostracodes, polychaetes, small 
gasteropods and bivalves 
Acanthomunna spinipes 
Anthurids 

Excirolana bicornis 

Gnathia sp. 
Haploniscus gernekei 
Ilyarachna wolffi 

Microarcturus youngi 

Neastacilla longispina 

Spinarcturus natalensis 
Stenetrium abyssale 
Stenetrium dalmeida 
Stenetrium dagama 



SM 103 

680 metres 

Light biological dredge 
Fine hard mud, overlain with hetero- 
pod and pteropod shells, coral 
fragments, numerous foraminifera, 
sponge fragments and spicules 
Hydroids, small echinoderms, pycno- 
gonids, amphipods, small crabs, 
small gasteropods and bivalves. 



Acanthomunna spinipes 
Anthurids 
Bathycopea typhlops 

Eurycope glabra 

Haploniscus gernekei 
Ilyarachna wolffi 
Ilyarachna sp. 
Microarcturus youngi 
Microarcturus oudops 

Serolis brinki 
Spinarcturus natalensis 
Stenetrium abyssale 

Stenetrium dagama 



The affinities of this fauna are not easily apparent, especially with 52 per 
cent being described as new. The presence of a Serolis, and Stenetrium abyssale 
and Acanthomunna spinipes indicate possible Antarctic/subantarctic affinities, 
while five endemic species {Microarcturus oudops, Stenetrium dagama, Cirolana 
imposita, and two easily recognizable anthurids Leptanthura laevigata and 
Mesanthura catenula) show the expected South African component. The 
unexpected presence of Bathycopea typhlops can only be explained by the 
paucity of collecting in this type of habitat, especially off the west African coast. 



THE SOUTH AFRICAN MUSEUM'S MEIRING NAUDE CRUISES 157 

ACKNOWLEDGEMENTS 

I should like to thank the crew of the R/V Meiring Naude and my colleagues 
of the South African Museum who assisted in the collecting of the material 
described here. I am grateful to Dr T. E. Bowman of the Smithsonian Institution, 
and Prof. J. H. Day of the University of Cape Town, who read the manuscript 
and made many useful criticisms and suggestions. 

REFERENCES 

Barnard, K. H. 1914. Contributions to the Crustacean fauna of South Africa. 3. Additions 

to the marine Isopoda, with notes on some previously incompletely known species. 

Ann. S. Afr. Mus. 10: 325-442. 
Barnard, K. H. 1920. Contributions to the Crustacean fauna of South Africa. 6. Further 

additions to the list of marine Isopoda. Ann. S. Afr. Mus. 17: 319-438. 
Barnard, K. H. 1940. Contributions to the Crustacean fauna of South Africa. 12. Further 

additions to the Tanaidacea, Isopoda and Amphipoda, together with keys for the 

identification of the hitherto recorded marine and freshwater species. Ann. S. Afr. Mus. 

32: 381-543. 
Bowman, T. E. 1971. Excirolana kumari, a new tubicolous isopod from Malaysia. Crustaceana 

20: 70-76. 
Hale, H. M. 1946. Isopoda -Valvifera. Rep. B.A.N.Z. antarct. Res. Exped. 4: 163-212. 
Hessler, R. R. 1970. A new species of Serolidae (Isopoda) from bathyal depths of the 

equatorial Atlantic Ocean. Crustaceana 18: 227-232. 
Kensley, B. 1975. Marine Isopoda from the continental shelf of South Africa. Ann. S. Afr. 

Mus. 67: 35-89. 
Louw, A. E. 1977. The South African Museum's Meiring Naude cruises. Part 1. Station Data 

1975, 1976. Ann. S. Afr. Mus. 72: 147-159. 
Loyola e Silva, J. 1971. Sobre os generos Ancinus Milne Edwards, 1840 e Bathycopea 

Tattersall, 1909, da colecao U.S. Nat. Mus. Archos Mus. nac, Rio de J. 54: 209-223. 
Menzies, R. J. 1962. The isopods of abyssal depths in the Atlantic Ocean. Vema Res. Ser. 1: 

84-206. 
Nordenstam, A. 1933. Marine Isopoda of the families Serolidae, Idotheidae, Pseudidotheidae, 

Arcturidae, Parasellidae, and Stenetriidae mainly from the South Atlantic. Further zool. 

Results Swed. Antarct. Exped. 3 (1): 1-284. 
Schultz, G. A. 1973. Ancinus H. Milne Edwards in the New World (Isopoda, Flabellifera). 

Crustaceana 25: 267-275. 
Tattersall, W. M. 1905. The marine fauna of the coast of Ireland. Part V. Isopoda. Scient. 

Invest. Fish, brch Ire. 1904 (2): 1-90. 
Vanhoffen, E. 1914. Dielsopoden derDeutschen siidpolar-Expedition 1901-1903. Dt. Siidpol.- 

Exped. 14: 447-598. 
Wolff, T. 1962. The systematics and biology of bathyal and abyssal Isopoda Asellota. 

Galathea Rep. 6: 1-320. 



6. SYSTEMATIC papers must conform to the International code of zoological nomenclature 
(particularly Articles 22 and 51). 

Names of new taxa, combinations, synonyms, etc., when used for the first time, must be 
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Family Nuculanidae 

Nuculana (Lembulus) bicuspidata (Gould, 1 845) 

Figs 14-1 5A 
Nitctila (Leda) bicuspidata Gould, 1845: 37. 
Leda plicifera A. Adams, 1856: 50. 

Laeda bicuspidata Hanley, 1859: 118, pi. 228 (fig. 73). Sowerby, 1871 : pi. 2 (fig. 8a-b). 
Nucula largillierti Philippi, 1861 : 87. 
Leda bicuspidata: Nickles, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9. 

Note punctuation in the above example: 

comma separates author's name and year 

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In describing new species, one specimen must be designated as the holotype; other speci- 
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not regarded as paratypes should be listed separately. The complete data (registration number, 
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Holotype 

SAM-A13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King's Beach 
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Name of new genus or species is not to be included in the title : it should be included in the 

abstract, counter to Recommendation 23 of the Code, to meet the requirements of 

Biological Abstracts. 



BRIAN KENSLEY 

THE SOUTH AFRICAN MUSEUM'S 
MEIRING NAUDE CRUISES 

PART 7 
MARINE ISOPODA 



VOLUME 74 PART 6 JANUARY 1978 



ISSN 0303-2515 



MUS, COMP. ZOOL 
LIBRARY 

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UNIVERSITY 



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(a) Author's name and year of publication given in text, e.g. : 

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within each name, with suffixes a, b, etc. to the year for more than one paper by the same 
author in that year, e.g. Smith (1969a, 1969Z>) and not Smith (1969, 1969a). 

For books give title in italics, edition, volume number, place of publication, publisher. 

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number (only if independently paged) in parentheses, pagination (first and last pages of article). 

Examples (note capitalization and punctuation) 

Bullough, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan. 

Fischer, P.-H. 1948. Donnees sur la resistance et de le vitalite des mollusques. /. Conch., Paris 88: 100-140. 

Fischer, P.-H., Duval, M. & Raffy, A. 1933. Etudes sur les echanges respiratoires des littorines. Archs 

Zool. exp. gen. 74: 627-634. 
Kohn, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon. 

Ann. Mag. nat. Hist. (13) 2: 309-320. 
Kohn, A. J. 19606. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean. 

Bull. Bingham oceanogr. Coll. 17 (4): 1-51. 
Twele, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. In: Schultze, L. Zoologische 

und anthropologische Ergebnisse einer Forschungsreise im west lichen und zentralen Sud-Afrika 4: 269-270. 

Jena: Fischer. Denkschr. med.-naturw. Ges. Jena 16: 269-270. 

(continued inside back cover) 



ANNALS OF THE SOUTH AFRICAN MUSEUM 
ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM 



Volume 74 Band 
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Part 6 Deel 




THE GEOGRAPHICAL DISTRIBUTION OF 
SOUTHERN AFRICAN HYDROIDS 



By 
N. A. H. MILLARD 



Cape Town 



Kaapstad 



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THE GEOGRAPHICAL DISTRIBUTION OF 
SOUTHERN AFRICAN HYDROIDS 

By 

N. A. H. Millard 

South African Museum, Cape Town 

(With 9 figures, 8 tables and 2 appendices) 

[MS. accepted 9 August 1977] 

ABSTRACT 

The geographical distribution of 251 species of hydroid Cnidaria round the coast of 
southern Africa is analysed by means of a system of radial sectors. It is concluded that there is 
no distinct west coast fauna as such, but that a boundary zone exists at approximately 31°S 
latitude between an east coast region with mainly tropical affinities and a temperate west-plus- 
south coast region with a reduced tropical component and a large endemic population. The 
main centre of the endemic population is the Agulhas Bank. The relationships of the tropical 
species are mainly with the Western Indian Ocean Province and the Indo-West-Pacific Region. 
Deep-water species (45) which occur below 400 m are of a mixed nature and show no clear 
relationship with other parts of the world. Deep-water species are classified into stenobathic/ 
eurybathic and stenothermic/eurythermic categories, and their distribution relative to water 
temperatures is discussed. 

CONTENTS 

PAGE 

Introduction 159 

Methods 160 

Faunal boundaries 164 

The tropical component 169 

The temperate component 170 

The endemic component 173 

The cosmopolitan and scattered components . . 174 

Distribution and depth 175 

Estuarine and brack-water species . . . . . 179 

Conclusion 182 

References 183 

Appendix 1 185 

Appendix 2 188 

INTRODUCTION 

This paper is an attempt to analyse the distribution of the hydroid fauna of 
the coast of southern Africa as documented in a recent monograph (Millard 
1975). Since the latter went to press a few more taxonomic papers on, or lists of, 
hydroids have appeared, namely Arnaud et al. (1976), Beurois (1975), Calder 
(1975, 1976), Cooke (1975), Cornelius (1975a, 19756), Mergner & Wedler 
(1977), Millard & Bouillon (1975), Millard (1977a) and Watson (1975). Among 

159 
Ann. S. Afr. Mm. 74 (6), 1978: 159-200, 9 figs, 8 tables, 2 appendices. 



160 ANNALS OF THE SOUTH AFRICAN MUSEUM 

these are records which extend the range of some South African species and they 
have been taken into account in the analysis which follows, as also have records 
from Buchanan (1957), Michel (1974), Blanco (1967, 1968, 1973, 1974) and 
Blanco & Bellusci de Miralles (1972a, 1972&), which have only recently become 
available to the author. 

Within the borders of southern Africa, here taken to be the 20° parallel of 
south latitude, several new distribution records from deep water off the Natal 
coast and some new records for the country were published by Millard (1977 b). 
There are also scattered identifications by the author, as yet unpublished, which 
are listed in Appendix 1 . 

The following changes in name have been adopted here : 
Antennella africana Broch, 1914, to A. quadriaurita Ritchie, \909, fide Millard 

(19776: 123). 
Eulaomedea caleeolifera (Hincks, 1871) to Campanularia calceolifera Hincks, 

\%l\,fide Cornelius (1975a: 254).* 
Lafoeafruticosa (M. Sars, 1851) to Lafoea dumosa (Fleming, \%2Q)fide Cornelius 

(19756: 385). 
Obelia bicuspidata Clarke, 1875, to O. bidentata Clarke, \815, fide Cornelius 

(1975a: 260). 

Any zoogeographical analysis is aimed at pinpointing population changes 
within an area, and, if possible, relating them to physical or chemical factors 
and determining relationships with other areas. Such an analysis is subject to 
certain unavoidable weaknesses— the assignment of species to categories or 
components is largely subjective; numbers of species within categories are 
liable to change as more records are forthcoming (for instance some of the 
South African endemics may in time be expected to turn up outside our 
boundaries); and numbers of species may be influenced by different intensities' 
of collecting in different areas or by misidentifications. The longer we wait the 
more reliable will be the results, but results, if only interim ones, are needed 
now. The larger the number of species involved the more meaningful will be the 
conclusions. One must bear in mind that it is the main trends which are 
significant rather than the details. 

METHODS 

The first step in the analysis was to remove all doubtful records and 
incomplete identifications. This left a total of 251 species. 

The first objective was an analysis of distribution within the borders of 
southern Africa, and for this some system of comparison was necessary. When 
dealing with the littoral zone it is a comparatively simple matter to convert the 
distance along the coast to a straight line, a method with obvious practical 

* Cornelius pointed out that the type species of Campanularia is Sertularia verticil/ata 
Linnaeus, 1758, and not Sertularia volubilis Linnaeus, 1758, as previously thought (Millard 
1975: 203). Since S. verticillata is a branched species the diagnosis of Campanularia given 
previously (Millard 1975: 203) must be altered to include branching forms, and Eulaomedea 
sunk in Campanularia. 






THE GEOGRAPHICAL DISTRIBUTION OF SOUTHERN AFRICAN HYDROIDS 



161 



CN 



CN 


CO 


■«* 


«o 


•o 


K 


00 


CO 


CO 


CO 


CO 


CO 


co 


co 




o — 

CN CN 



TT 


«n 


-o 


IS. 


00 


O 


O 


CM 


CN 


CN 


CN 


CN 


CN 


CO 



CM 


CO 


"* 


m 


•o 


K 


00 


co 


CO 


CO 


CO 


CO 


CO 


CO 



162 



ANNALS OF THE SOUTH AFRICAN MUSEUM 



advantages, and this has been done by Stephenson (1947), Stephenson & 
Stephenson (1972), Day (1967) and others. However, when dealing with the sub- 
littoral area as well, this is clearly not possible for a continent of the shape of 
southern Africa. After much consideration the following arbitrary method was 
devised, in which the coastal waters were divided into thirty-six sectors of 
approximately equal area (Fig. 2). 

The central point for the sectors was chosen as the intersection of the 
latitude line of 20°S and the longitude line of 24°E, a point which is approxi- 
mately mid-way between the west and east coastlines. From this central point 
radii were drawn at 5° intervals, starting with one through the Cape Peninsula, 
and then working outwards in both directions. The sectors thus constructed 
were numbered 1-36. The reason for placing one radius (Number 14/15) 
through the Cape Peninsula is that Cape Point is generally recognized as an 
approximate zoogeographical boundary between the cold Benguela water on 
the west and the warmer Agulhas water on the east, and might be expected to 
have some significance. It so happens that another radius (Number 15/16) 
conveniently passes through Cape Agulhas, the most southerly point of Africa, 
and a third (Number 23/24) through the boundary between Transkei and Natal. 




Fig. 2. Map of southern Africa illustrating the sector system which was used to analyse the 

distribution. 



THE GEOGRAPHICAL DISTRIBUTION OF SOUTHERN AFRICAN HYDROIDS 163 

It must be emphasized that this is an arbitrary method, and obviously 
could not be applied to all coastlines, but it is felt that it might form a useful 
working method and basis for comparison with other southern African marine 
groups. It has the advantage that each sector covers an approximately equal 
area of sea and 'straight-line' length of coast and represents a cross-section 
through all depths. Since the sectors are roughly normal to the coast and the 
currents tend to follow the coast, each sector is affected in a similar way and 
population differences related to water temperature should be highlighted. It is 
felt that the advantages of this method at least outweigh the disadvantages (and 
impracticabilities) of the 'straightening-out' method of Stephenson. 

The second objective was an analysis of the world-wide distribution of the 
southern African species, and for this the species were delegated to components 
defined as follows: 

TROPICAL 

Circumtropical. Present in the tropics of all three oceans, but may spread 
north or south into warm temperate seas as well. 

Indo-West-Pacific Region. Stretching from the Suez Canal across the Indian 
Ocean and the East Indies into the Western Pacific as far as Hawaii and 
Easter Island. Includes northern Australia, Madagascar and East Africa. 
May spread into warm temperate seas as well. 

Western Indian Ocean Province. The western section of the Indo-West- 
Pacific Region, bounded in the north by the Persian Gulf, and including 
Madagascar and surrounding islands. 

temperate. (In a trial analysis it was found impossible to differentiate between 
cold temperate and warm temperate fauna when assessing world distribution.) 

Antitropical {bipolar). Present in the temperate zones of all three oceans, 

and present in both the Northern and the Southern hemisphere, but not 

in the tropics except perhaps just at the edge. 

Southern. Restricted to the temperate waters of the Southern hemisphere, 

but often in the subantarctic islands as well, and rarely reaching the 

Antarctic. 

Atlantic. Restricted to the northern and/or southern temperate zones of 

the Atlantic, but absent from the tropics. May spread further north into 

the subarctic. 

ENDEMIC 
OTHER 

Cosmopolitan. Present in all three oceans, and occurring in both tropical 
and temperate seas. 

Scattered. Present in various regions, but not sufficiently widespread to 
be included in Cosmopolitan. Does not fit any other category. 



164 ANNALS OF THE SOUTH AFRICAN MUSEUM 

A list of species, together with their components and distribution within 
thirty-four sectors, is given in Appendix 2. (Sectors 35 and 36 have been omitted 
as no records were available from this region.) 

FAUNAL BOUNDARIES 

It has long been recognized that the marine fauna of southern Africa 
includes both tropical elements, predominating on the east coast, and temperate 
elements, predominating on the south and west coasts. Most authorities, 
following Stephenson (1947) and Stephenson & Stephenson (1972), have 
recognized a separate east coast (tropical or subtropical) fauna, a south coast 
fauna (warm temperate) and a west coast fauna (warm or cold temperate) for 
the littoral region. These earlier views are summarized by Briggs (1974), who 
includes the east coast in the Western Indian Ocean Province of the Indo-West- 
Pacific tropical region; the south coast in the Agulhas Province of the South 
African warm temperate region; and the west coast as a separate South West 
Africa Province of the South African warm temperate region. Opinions have 
differed as to the position of the boundary between the east and south coast, 
and whether the west coast represents a separate faunal province, and if so 
whether it should be regarded as cold temperate or warm temperate. 

The distribution of the faunal components of the continental shelf and the 
extent of the faunal regions is largely dependent on the ocean currents round 
the South African coast, in general the warm southward-flowing Agulhas 
Current being responsible for the southerly extension of tropical forms on the 
east coast, and the cold northward-flowing Benguela Current being responsible 
for the more northerly distribution of cool-water forms on the west coast. It is 
not intended to discuss these current systems in any detail here since accounts 
are available in most textbooks of marine biology (a concise general account is 
given by Stephenson & Stephenson (1972) and more recent contributions are 
summarized by M. J. Penrith (1976)). 

In order to determine whether there is any region of the coast where a 
marked change in the hydroid fauna occurs, the range of the tropical, temperate 
and endemic components was plotted separately (Fig. 3). (The range of species 
was used in preference to presence/absence, in order to minimize differences in 
collecting intensity.) This figure emphasizes the tropical nature of the east coast 
fauna, for there is a strong tropical component at the northern end which 
gradually drops away, until beyond St Helena Bay on the west coast (Sector 14) 
only one species remains for a short distance. The rate of disappearance is 
greatest at the boundary of Sectors 23/24, and is also high in the region of 
Sectors 13/14/15. The temperate and endemic components are both highest on 
the south coast, falling away fairly evenly to the north-east, but more sharply to 
the north-west in the region of Sectors 13/14/15. A similar trend appears when 
the range of species is plotted as a percentage of the total range (Fig. 4). 

It is thus possible to distinguish two 'zones of change', though they are not 
as marked as might be expected. The first (Sectors 23/24) provides a convenient 



THE GEOGRAPHICAL DISTRIBUTION OF SOUTHERN AFRICAN HYDROIDS 



165 



o 

LU 
Q_ 



20 i 
10 



50 J 

40 

30 

</> 20 



10 



O 

LU 

m 80 

3 



70 

60- 

50- 

40 

30 

20- 

10- 



(0 
00 

w 

i 



>» 

(0 
CO 

M 



0) 

3 



>» 




<o 


w 


CO 


CO 




+-.£ 


(0 


-E-5 


■»-> 


O O) 


2 

E 


Q_< 

CD CO 


Q.Q. 



CD 
XJ 
(0 
N 



c 
o 
■o 

c 
o 



J 1 

TEMPERATE 



{0 CO to 

JUU 

HI 



O CO 

Q_ UJ 



c 

SI 

I— 
■3 

Q 



(0 

o 

(0 



1 1 1 




ENDEMIC 




TROPICAL 



1 



10 




15 20 

SECTORS 



Fig. 3. The distribution of the temperate component (above), the endemic component (centre) 
and the tropical component (below). From range of species. 



166 



ANNALS OF THE SOUTH AFRICAN MUSEUM 



boundary between the east and south coast fauna. It is located between 30 and 
32°S latitude, i.e. the stretch of coast between the Bashee River in the south and 
Durban in the north. For the discussions which follow, the boundary between 
Sectors 23 and 24 is used (31°S). This is more or less in agreement with con- 
clusions drawn from other faunal groups. Stephenson & Stephenson (1972) 
suggest a boundary zone between Port St Johns and Qolora, i.e. at about 32°S, 
for the intertidal fauna (the form of the graph of the tropical component in 
Figure 3 of this paper is very similar to that for the intertidal fauna in 
Stephenson & Stephenson (1972, fig. 8.26)); J. L. B. Smith (1949) and 
M.-L. Penrith (1970) suggest the Great Kei River at 32°42'S for the fishes, and 
Day (1967) suggests the Bashee River at 32°15'S for the benthic Polychaeta. 



TEMPERATE 




S ECTORS 

Fig. 4. The distribution of the temperate, endemic and tropical components, from range of 

species as a percentage of the total range. Sectors 1-7 have been omitted since the total number 

of species is less than twenty and the percentages unrealistic. 



THE GEOGRAPHICAL DISTRIBUTION OF SOUTHERN AFRICAN HYDROIDS 



167 



At the boundary of Sectors 23/24 the monthly minimum surface temperature 
near the coast does not fall below 16°C, the maximum does not rise above 27°C 
and the mean ranges from 21 to 24°C (U.S. Naval Oceanographic Office 1967). 
This is approximately the region where the Agulhas Bank begins to widen and 
force the warm Agulhas Current offshore. Further south the latter is separated 
from the coast by a cooler counter-current flowing northwards. 

The usefulness of the second 'zone of change' in the south-westerly corner 
of the Cape Province (Sectors 13/14/15) is more debatable. It could be used to 
divide the temperate region, which stretches from north of 20°S on the west 
coast to 30-32°S on the east coast, into two provinces, if warranted by the 
fauna. 

To test this an arbitrary boundary was placed at Cape Point (Sector 14/15) 
and the composition of species on the two sides of it compared with one another 
and with the east coast (Table 1). (Cape Point was used as a boundary by 
Stephenson (1947, fig. 3), Ekman (1953) and Briggs (1974).) It may be noted 
from Figures 3 and 4 that there is no evidence of a change in fauna at Cape 
Agulhas (Sector 15/16), the most southerly point of Africa). 



Table 1 

Analysis of fauna of west coast, as compared with that of the south and east coasts 

(actual records, not range). 



Component 



West coast 
(Sectors 1-14) 



No. of 
species 



South coast 
(Sectors 15-23) 



East coast 
(Sectors 24-36) 



No. of 
species 



No. of 
species 



Tropical 

Temperate . 

Endemic to area 

Endemic to more than one 

Cosmopolitan 

Scattered 



Total . 



area 



13 

2 
25 
17 
18 



9,6 

15,7 
2,4 
30,1 
20,5 
21,7 



30 
20 

21 
41 
21 

25 



19,0 
12,7 
13,3 
25,9 
13,3 
15,8 



72 
12 
17 
30 
28 
23 



83 



100,0 



158 



100,0 



182 



39,6 
6,6 
9,3 
16,5 
15,4 
12,6 



100,0 



It is apparent from Table 1 that the west coast fauna consists mainly of 
endemic, cosmopolitan and scattered species. The tropical species diminish in 
number round the coast from east to west; only 8 species remain on the west 
coast and these are all absent north of St Helena Bay. The temperate species of 
the west coast are fewer in number than those of the south coast. Only 2 of them 
{Clavopsella navis and Gonothyraea loveni, both from Table Bay harbour and 
probably introduced by ships), and possibly a third (Sarsia eximia), are 
restricted to the west coast. Only 2 rare species are endemic solely to the west 
coast (Myriothela tentaculata and Eudendrium ritchiei), whereas 21 are endemic 
to the south coast and 17 to the east coast. Of the cosmopolitan and scattered 
components only 3 (Acryptolaria crassicaulis, Hybocodon unicus and Hydro- 
dendron gracilis) have been recorded from the west coast alone. 



168 ANNALS OF THE SOUTH AFRICAN MUSEUM 

In total, thus, the west coast has only 8 unique species (9,8%), of which 
only 2 (2,4%) are endemic to the area. Briggs (1974) requires at least 10 per cent 
endemic species for the designation of a separate province, and on this basis 
the west coast of South Africa does not qualify, at least for the hydroids. The 
hydroid fauna of the west coast seems to be mainly an impoverished south coast 
fauna. 

Although Day (1967) accepted a separate west coast element for the littoral 
polychaets, he could find no marked difference between the south and west 
coasts for the sublittoral ones, and ascribed this to the fairly uniform water 
temperature on the bottom of the continental shelf (he gave figures of 12-14°C 
at 100 m from Port Elizabeth to Liideritz, but the records collated by Hulley 
(1972) show a somewhat lower range of 8-1 3°C at 100 m between 20°S on the 
west coast and Port Elizabeth on the south coast). On the other hand there is a 
difference in the surface water temperature on the two sides of Cape Point 
(Day gives a range of 15-20°C for the south coast and 12-1 5°C for the west 
coast, but under certain conditions and for certain periods the south coast 
temperature may drop to as low as 11°C (during local upwelling) and rise as 
high as 24°C, while that of the west coast may drop to 9°C (M. J. Penrith 1976)). 
In this respect it might be noted that all hydroids are sublittoral in the sense 
that they cannot withstand drying and survive in the littoral zone only in damp 
overhangs or open runnels. 

None of this really explains why there should be a diminution of hydroid 
species on the west coast and practically no endemic population of cold- 
temperate forms. Ekman (1953) associates the poverty of the benthos of the 
'Namaqua coast' with the hydrogen sulphide resulting from frequent upwelling 
and red tides. Briggs (1974) suggests that certain species are prevented from 
spreading westwards round Cape Point by the absence of high summer 
temperatures for reproduction (he gives a February mean of 16-18°C for South 
West Africa as against one of 20-22°C for the south coast, but compare remarks 
in parenthesis above). M.-L. Penrith & Kensley (1970a) suggest that a region of 
minimal temperatures between Hondeklip Bay and Liideritz (i.e. north of 
St Helena Bay) acts as a barrier to the establishment of southern warm- 
temperate intertidal species further north. The paucity of hydroid records on the 
west coast may be partly a reflection of the difficulty of collecting on the exposed 
and inhospitable Skeleton Coast. However, M.-L. Penrith (pers. comm.), who 
has collected in this area, feels that the poverty is a real one, and perhaps linked 
to the absence of suitable habitats such as sheltered coves, rock pools, etc., and 
the preponderance of surf beaches and sandy floors. The harsh conditions and 
poverty of the rocky-shore fauna are also emphasized by M. J. Penrith (1976). 

There is no evidence at all of a southward spread of species from tropical 
west Africa. The southern boundary of the west African tropical region is too 
far north for consideration in this paper (14 or 15°S: Briggs 1974; 18-22°S: 
Hulley 1972). M.-L. Penrith & Kensley (19706) found the first of the west 
African littoral species at Rocky Point (18°59'S). 



THE GEOGRAPHICAL DISTRIBUTION OF SOUTHERN AFRICAN HYDROIDS 169 

In conclusion the south and west coast sublittoral areas are considered to 
represent a single temperate province for the hydroid fauna, which might be 
termed the Agulhas/Namaqua Province, the centre of the endemic population 
being on the Agulhas Bank. 

THE TROPICAL COMPONENT 

Of the total number of 251 species, 77 (30,7%) can be classified as tropical. 
Of these, 20 are circumtropical, 24 Indo-West-Pacinc and 33 confined to the 
Western Indian Ocean. In addition there are 17 endemic species which are 
confined to the South African east coast, and therefore with tropical tendencies 
(these were termed 'subtropical endemics' by Day (1974)). (Appendix 2.) 

Ekman (1953) and Briggs (1974) consider that the main centre of origin of 
the tropical fauna is the Indo-Malayan Region. From here species tend to 
spread west to the Indian Ocean (and may penetrate into the Red Sea or round 
southern Africa), and to a lesser extent east to the Eastern Pacific. 

Species have been classified here as Indo-West-Pacific or Western Indian 
Ocean even though a few have one or two records outside these areas. Thus, 
2 have penetrated into the Mediterranean {Corydendrium parasiticum, Thyro- 
scyphus fruticosus), 4 have reached the tropical west coast of Africa (Abietinaria 
laevimarginata, Campanularia africana, Corydendrium parasiticum, Sertularia 
ligulata), 1 has reached tropical South America {Dentitheca bidentata), and 1 has 
reached southern Australia {Dynamena obliqua). Further spread would produce 
a circumtropical distribution, but of the species classified as such no less than 
13 have failed to reach the east coast of the Pacific, and this applies also to 4 of 
the cosmopolitans (Table 2). There is therefore support for Briggs's contention 
(1974) that the 'East Pacific barrier' is one not easily crossed. 

There is some evidence for a secondary centre of origin in the Western 
Indian Ocean, for in addition to the seventeen South African subtropical 
endemics (i.e. present on the east coast only), there are seven Western Indian 

Table 2 

List of circumtropical and cosmopolitan species which 
do not occur on the east coast of the Pacific. 

Circumtropical Cosmopolitan 



Diphasia digitalis Antennella secundaria 

Dynamena quadridentata Cladocoryne floccosa 

Gymnangium hians Coryne pusilla 

Halecium dyssymetrum Halecium sessile 

Hydrodendron caeiniformis 

Jdiellana prist is 

Monostaechas quadridens 

Plumularia strictoearpa 

Scandia mutabilis 

Sertularella diaphana 

Sertularia distans 

S. marginata 

S. turbinata 



170 ANNALS OF THE SOUTH AFRICAN MUSEUM 

Ocean species which, though absent from the Indo-Polynesian area, have yet 
penetrated into the Red Sea (Clytia latitheca, Cytaeis nassa, Diphasia heurteli, 
Halopteris glutinosa, Hydractini kaffraria, Plumularia wasini, Solanderia 
minima). 

Some communication between southern Australia and the Western Indian 
Ocean is indicated, since there are five species common to the two, namely 
Amphisbetia maplestonei, Crateritheca acantho stoma, Halopteris glutinosa, 
Sertularella arbuscula and Synthecium dentigerum. These occur nowhere else 
(except for H. glutinosa which also occurs in the Red Sea) and the likelihood is 
that the movement has been from west to east with the West Wind Drift. 

THE TEMPERATE COMPONENT 

The total temperate component includes 28 species (11,2%), of which 
11 are antitropical (bipolar), 10 are restricted to the Southern hemisphere and 
7 are restricted to the Atlantic (Appendix 2). These are species which cannot 
survive the high temperatures of the tropics, although some can spread into the 
cooler waters of the subpolar regions. The southern species must have evolved 
in the south, but the antitropical and Atlantic species may have evolved in 
either the south or the north and must at some time have crossed the tropics. 
This may have been achieved by equatorial submergence, for Briggs (1974) 
points out that the tropical surface water is only about 30-40 m deep in the 
Eastern Atlantic. It may also be due to transportation on the hulls of ships. 
In all, twenty-one species have been recorded on ships' hulls or floating objects 
such as buoys. The fauna of ships' hulls is a characteristic group of hardy 
species (Millard 1952), most of them cosmopolitan. Three of the species (all 
belonging to the temperate component) are known only in harbour areas 
in South Africa, and it is suggested that these are recent introductions by 
ships : 

Gonothyraea loveni is well established in Cape Town docks, occurring 
abundantly on pylons, cables, etc., as well as on hulls of ships. It has not been 
reported from any other part of South Africa. Elsewhere it is known from the 
North Atlantic, New Zealand, Tasmania and the Argentine. 

Campanularia calceolifera has been found, and reproducing, in Cape Town 
docks on a barge and on a raft, neither of which had left the harbour. It must, 
therefore, be established, though not common. It is not known for certain 
elsewhere in South Africa, a record from False Bay by Stechow (1925) being of 
infertile material and thus subject to doubt. The species is well known on 
European and Mediterranean coasts and also occurs on the east coast of North 
America. 

Clavopsella navis is known from a ship's hull in Cape Town docks, and also 
from the Kiel Canal (Thiel 1962 as C. quadranularia). The latter is probably the 
original locality since the colonies there were more abundant and better 
developed. 



THE GEOGRAPHICAL DISTRIBUTION OF SOUTHERN AFRICAN HYDROIDS 



171 



Since all these three species occur both in the South and North Atlantic 
they must have traversed the tropics, though they have not established them- 
selves in tropical latitudes so far as is known. They may have been hardy 
enough to survive a brief passage through warmer waters on ships' hulls, or 
they may have died back during transport, leaving the hydrorhiza to regenerate 
again in suitable temperature conditions. Ralph (1961) also found evidence of 
the introduction of hydroids by ships' hulls to New Zealand harbours. 

It is interesting to compare the relations of the temperate species with the 
other two southern continents. From Table 3 it can be seen that South Africa 
has 9 species in common with southern Australasia alone, but only 3 species in 
common with South America alone. There are 7 species common to all three. 



Table 3 
Temperate species common to the southern continents. 



South Africa and South 
Australasia 



South Africa and temperate 
South America 



Temperate part of all 

three southern 

continents 



Kirchenpaueria triangulata 
Nemertesia ciliata 
Parascyphus simplex 
Plumularia obliqua 
P. spinulosa 
Sarsia eximia 

Sertularella annulaventricosa 
Stereotheca elongata 
Symplectoscyphus macrogonus 

(Total: 9) 



Phialella turrita 
Sertularella striata 
Symplectoscyphus paulensis 



(Total: 3) 



Amphisbetia minima 
Filellum antarcticum 
Gonothyraea loveni 
Nemertesia cymodocea 
Plumularia filicaulis 
Staurocladia vallentini 
Tubularia larynx 



(Total: 7) 



This would suggest dispersal eastward from the Agulhas Bank. Beurois (1975) 
emphasized the role of eastward drifting kelp in the dispersal of attached forms 
to St Paul and Amsterdam Islands, and Arnaud et al. (1976) discussed the 
transport of South African fauna to St Helena on kelp. If the list in Table 3 is 
extended to include the cosmopolitan and scattered components and those 
tropical forms which have spread south of the Tropic of Capricorn, the relation- 
ship between South Africa and Australasia is more marked (34 species in 
common with Australasia alone, 10 with South America alone, and 26 in all 
three). 

The distribution of those South African species which have antarctic or 
subantarctic affinities is given in Table 4. In all, 41 species are listed, a relatively 
small number (16,3% of the total), and of these half (21) are cosmopolitan and 
this tends to mask the relationships. Without the cosmopolitan species only 20 
(8,0%) have antarctic affinities and of these 11 are classified as temperate. The 
absence from South Africa of certain common antarctic genera, such as 
Stawotheca, Tulpa, Grammaria and Silicularia, is very striking. On the whole 
there is very little evidence of relationship with the Antarctic. 



172 



ANNALS OF THE SOUTH AFRICAN MUSEUM 



Table 4 

Distribution of species with antarctic/subantarctic affinities. The Magellan Region includes 
Gauss Station, the Falklands and South Georgia. The Kerguelen Group includes Marion, 

Prince Edward and Crozet Islands. 



Species 


Component 


.5 x; 

C 3 

o o 


c 
o 

'5b 

(L> 


Oh 

o 

S-H 


O 

u 

C 








\^ Of) 


_o3 
00 




13 

3 


3 






"8*2 


a 


_co 




Ph 








2 




^ 


Xfl 


Acryptolaria confer t a 


Cosmopolitan 


X 


X 








A. crassicaulis 


Scattered 


X 


X 








Aglaophenia p. pluma 


Cosmopolitan 




X 






X 


Amphisbetia minima 


Temperate (Antitr.) 




X 






X 


A. operculata 


Cosmopolitan 




X 




X 


X 


Antennella quadriaurita 


Scattered 






X 






A. secundaria 


Cosmopolitan 










X 


Bimeria vestita 


Cosmopolitan 




X 








Bougainvillia macloviana 


Scattered 




X 




X 




Campanularia crenata 


Cosmopolitan 




X 








C. integra 


Cosmopolitan 




X 








Clytia hemisphaerica 


Cosmopolitan 




X 






X 


C. paulensis 


Scattered 


X 








X 


Coryne pusilla 


Cosmopolitan 








X 




Dynamena cornicina 


Cosmopolitan 










X 


Filellum antarcticum 


Temperate (Southern) 


X 


X 








F. serratum 


Cosmopolitan 




X 




X 




Halecium beanii 


Cosmopolitan 




X 








H. delicatulum 


Cosmopolitan 


X 


X 




X 




H. tenellum 


Cosmopolitan 


X 


X 


X 


X 




Hincksella echinocarpa 


Temperate (Southern) 








X 




Hybocodon unicus 


Scattered 




X 








Kirchenpaueria triangulata 


Temperate (Southern) 








X 




Lafoea benthophila 


Scattered 


X 


X 




X 


X 


L. dumosa 


Cosmopolitan 


X 


X 




X 




Modeeria rotunda 


Cosmopolitan 




X 




X 




Nemertesia cymodocea 


Temperate (Southern) 




X 








N. ramosa 


Scattered 




X 








Obelia bidentata 


Cosmopolitan 




X 


X 


X 




0. dichotoma 


Cosmopolitan 


X 


X 


X 




X 


O. geniculata 


Cosmopolitan 




X 


X 


X 


X 


Parascyphus simplex 


Temperate (Antitr.) 


X 




X 






Phialella turrita 


Temperate (Antitr.) 




X 








Plumularia pulchella 


Scattered 




X 








P. setacea 


Cosmopolitan 




X 






X 


Sertularella leiocarpa 


Temperate (Southern) 






X 




X 


S. p. polyzonias 


Cosmopolitan 




X 




X 




S. striata 


Temperate (Southern) 




X 








Staurocladia vallentini 


Temperate (Southern) 




X 




X- 




Symplectoscyphus paulensis 


Temperate (Southern) 




X 






X 


Zygophylax armata 


Scattered 






X 







THE GEOGRAPHICAL DISTRIBUTION OF SOUTHERN AFRICAN HYDROIDS 173 

THE ENDEMIC COMPONENT 

The number of endemic species strictly belonging to the South African 
coast south of 20°S is 76. However, there are 2 species which also occur 
immediately north of 20° on the west coast (Gattya humilis and Salacia 
articulata). 4 species occur only in South Africa and the Vema Seamount 
{Halopteris pseudoconstricta, Salacia articulata, Sertularella flabellum and 
S. megista), and it was shown by Millard (1966) that the hydroid fauna of the 
Vema Seamount is essentially South African (21 of the 23 species also occur in 
South Africa). If these 6 species are included, the endemics number 82 (32,7% 
of the total of 251). It might be noted, however, that Mergner & Wedler (1977) 
have recorded Eudendrium deciduum and Sertularella natalensis from the 
Red Sea with a query. 

As a comparison the following percentages of endemics have been given 
for other large marine groups in southern Africa. The figures are not strictly 
comparable, however, since the boundaries used for the 'Southern African 
Region' are not always the same. 

Polychaeta 36% (Day 1967) 

Echinoderms, other than Holothuria 48% (Clark & Courtman-Stock 1976) 

Ascidians c. 50% (Millar 1971) 

Fish 25,3% (M. M. Smith 1970: 380 out of 

1 500 species) 

The south coast of southern Africa has more endemic species of hydroid 
than any other region, and the Agulhas Bank seems to be the centre of distri- 
bution of the endemic population (Figs 3-5). 

There are only three genera endemic to South Africa, Bicorona, Hydro- 
corella and Uniscyphus, and all of these are monotypic. However, certain genera 
have a larger proportion of endemic species than others (Table 5). These are 
probably genera in which speciation is actively occurring in the South African 
area. They are also genera in which many of the species are not easily dis- 
tinguished from one another, and thus pose constant problems to the systematist. 
Similarly, species with several subspecies are ones likely to be undergoing active 
speciation. Thus, Thecocarpus flexuosus has four subspecies; the nominal one is 

Table 5 

List of South African hydroid genera with the largest number of endemic species and/or 

subspecies. 

Number of endemic species Total number of species 

Genus or subspecies or subspecies in South Africa 

Cladocarpus 9 13 

Corhiza 5 5 

Halopteris 4 6 

Hydractinia 3 5 

Sertularella 14 23 

Zygophylax 4 7 



174 



ANNALS OF THE SOUTH AFRICAN MUSEUM 



WEST 



SOUTH 



EAST 




10 



15 20 

SECTORS 



Fig. 5. An analysis of the distribution of the endemic species, with the total number of species 
distributed between six components, as in Stephenson (1972). From range of species. 

tropical Indo-West-Pacific and the other three are endemic to southern Africa. 
Sertularella polyzonias also has four subspecies; the nominal one is cosmopolitan 
and two of the others are endemic to southern Africa. 

Several common genera, on the other hand, have no southern African 
endemics: Aglaophenia, Amphisbetia, Dynamena, Lytocarpus, Nemertesia, 
Obelia and Sertularia. 



THE COSMOPOLITAN AND SCATTERED COMPONENTS 

The cosmopolitan species number 28 (11,2%). They owe their wide 
distribution to the fact that they are eurythermic, hardy and adaptable. The 
wide distribution possibly indicates a very old history, in which there has been 
plenty of time to disperse. 

Species classified as scattered or disjunct number 36 (14,3 %). Most of them 



THE GEOGRAPHICAL DISTRIBUTION OF SOUTHERN AFRICAN HYDROIDS 



175 



have odd distributions which do not fit any of the other categories. There are 
two possibilities. Either these are newly evolved eurythermic species which are 
in the process of spreading rapidly and likely to become cosmopolitan, or they 
are 'peripheral relicts' of fairly primitive forms which were once more widely 
distributed. 

Among the scattered species three are interesting because outside southern 
Africa they are limited to the west and east coasts of the tropical Atlantic 
(Clytia hummelincki, Zyzzyzus solitarius) or to the western tropical Atlantic 
only {Cladocarpus tenuis). Three others have a similar distribution but also 
extend further north and/or east (Aglaophenia latecarinata, A. pluma dichotoma, 
Symplectoscyphus amphoriferus). 

Cosmopolitan and scattered species with antarctic/subantarctic affinities 
are included in Table 4. 

DISTRIBUTION AND DEPTH 

Figure 6 shows that there are more species present in the shallow water 
(0-100 m) than at any other level, and although there have been fewer samples 
from deep water it is not expected that more extensive collecting will alter the 
picture to any great extent. The shallow water is obviously the optimum zone 
for hydroids. Most of the species in the littoral area are at the upper edge of 



180 -i 




750 1000 1250 

DEPTH (m) 
Fig. 6. The distribution of species according to depth. From range of species 



1500 



176 ANNALS OF THE SOUTH AFRICAN MUSEUM 

their depth range and able to survive only under overhangs and in rock pools 
and crevices. Only 24 of the 98 littoral species have not been found below low 
water of springs. 

Deep-water species are those which occur over the edge of the continental 
shelf, and Ekman (1953) places an arbitrary upper boundary for the deep-sea 
fauna at 400 m. The number of hydroids known from below this depth round 
the South African coast is relatively small (45 = 1 7,9 %), and only eight of these 
reach 1 000 m (abyssal). The species are listed in Table 6. 

These species can be divided into two arbitrary categories on the basis of 
their depth range both in South Africa and the rest of the world : 
Stenobathic : those that are restricted to water deeper than 200 m. 
Eurybathic: those that occur above and below 200 m. 

These two categories are distributed between the different geographical 
components as follows : 

Stenobathic Eurybathic Total 

Tropical 5 5 10 

Temperate 1 4 5 

Endemic 5 6 11 

Cosmopolitan 9 9 

Scattered 1 9 10 

Total 12 33 45 

All the (12) stenobathic species are in reality cold-water species frequenting 
water below about 16°C (Ekman 1953, table 42), and this includes five so-called 
'tropical' species. These species are thus also stenothermic. Their limiting factor 
for distribution may be depth or temperature or a combination of both. 

Some of the eurybathic species are also stenothermic. This applies to the 
following tropical and scattered species whose records within the tropics are 
never shallow, and which are thus restricted to cold water: 

Most shallow record in tropics (m) 

Acryptolaria rectangularis (tropical) 228 

Cladocarpus tenuis (scattered) 185 

Cryptolarella abyssicola (scattered) 4 560 

Symplectoscyphus amphoriferus (scattered) 161 

Zygophylax sibogae (scattered) Only one tropical record : 

depth not given. 

To the same category of eurybathic/stenothermic species might be added 
the endemic and temperate eurybathic species, at any rate those restricted to the 
south-west coast of South Africa and the east coast below 200 m : 

Cladocarpus sinuosus (endemic): minimum east coast record 400 m. 
Hincksella echinocarpa (temperate) : minimum record Kerguelen 41 m, minimum 
east coast record 1 610 m. 



THE GEOGRAPHICAL DISTRIBUTION OF SOUTHERN AFRICAN HYDROIDS 



177 



Table 6 

South African deep-water species. 
SB— stenobathic (never recorded above 200 m); EB— eurybathic (occurring at all depths); 
ST— stenothermic (cold-water species); ET—eury thermic (occurring at all temperatures). 







Depth range 


Depth range 


Component 


Species 


Component 


in South 


outside South 


of depth J 






Africa (m) 


Africa (m) 


temperature 


Acryptolaria conferta 


Cosmopolitan 


64-1200 


0-4400 


EB/ET 


A. crassicaulis 


Scattered 


2835 


109-767 


EB/ET 


A. rectangularis ... 


Tropical 


110-1200 


228-600 


EB/ST 


Antennella quadriaurita 


Scattered 


0-825 


0-400 


EB/ET 


Branchiocerianthus imperator 


Tropical 


730 


200-5290 


SB/ST 


Campanularia hincksii 


Cosmopolitan 


9-450 


0-811 


EB/ET 


Cladocarpus distomus . 


Tropical 


292-2200 


55-1362 


EB/ET 




Tropical 


425-550 


1019 


SB/ST 






Endemic 


495 


— 


SB/ST 


C. millardae 




Tropical 


200-595 


5020 


SB/ST 


C. natalensis 




Endemic 


400-900 


— 


SB/ST 


C. sinuosus . 




Endemic 


183-680 


— 


EB/ST 






Scattered 


495 


185-550 


EB/ST 


Clytia gravieri 




Tropical 


0-528 


0-4243 


EB/ET 


Cryptolarella abyssicola 


Scattered 


100-2740 


745-4970 


EB/ST 


Filellum serratum 


Cosmopolitan 


24-900 


0-650 


EB/ET 


Garveia crassa 


Tropical 


625-900 


741 


SB/ST 


Halecium beanii . 


Cosmopolitan 


0-780 


0-1134 


EB/ET 


H. sessile .... 


Cosmopolitan 


425-430 


0-752 


EB/ET 


H. tenellum .... 


Cosmopolitan 


0-900 


0-730 


EB/ET 


Halopteris glutinosa . 


Tropical 


0-550 


1-80 


EB/ET 


H. polymorpha . 


Tropical 


0-900 


0-89 


EB/ET 


H. tuba .... 


Endemic 


11-550 


— 


EB/ET 


Hincksella echinocarpa 


Temperate 


1610-2200 


41-109 


EB/ST 


Hydrocorella africana 


Endemic 


0-500 


— 


EB/ET 


Kirchenpaueria triangulata 


Temperate 


111-1207 


550-585 


EB/ST 


Lafoea benthophila 


Scattered 


425-430 


672-3237 


SB/ST 


L. dumosa .... 


Cosmopolitan 


60-920 


0-2200 


EB/ET 


Modeeria rotunda 


Cosmopolitan 


70-550 


0-1240 


EB/ET 


Nemertesia antennina 


Scattered 


425-430 


0-1779 


EB/ET 


N. ramosa .... 


Scattered 


1 1-700 


0-872 


EB/ET 


Plumularia antonbruuni 


Endemic 


440 


— 


SB/ST 


P. mossambicae . 


Endemic 


110-550 


— 


EB/ET 


P. setacea .... 


Cosmopolitan 


0-430 


0-752 


EB/ET 


Sertularella leiocarpa 


Temperate 


200-900 


183-672 


EB/ST 


Stegolaria geniculata . 


Tropical 


775-850 


253-910 


SB/ST 


Symplectoscyphus amphoriferus 


Scattered 


550-900 


161-1256 


EB/ST 


S. arboriformis . 


Endemic 


10-420 


— 


EB/ET 


S. paulensis .... 


Temperate 


347-1200 


400-672 


SB/ST 


Uniscyphus fragilis 


Endemic 


360-420 


— 


SB/ST 


Zygophylax africana . 


Endemic 


137-850 


— 


EB/ST 


Z. armata .... 


Scattered 


48-440 


183 


EB/ET 




Temperate 


400-550 


20-752 


EB/ET 


Z. inconstans 


Endemic 


360-450 


— 


SB/ST 


Z. sibogae .... 


Scattered 


88-900 


146-550 


EB/ST 


Sui 


nmary : Cosmopol 


tan . 


9 






Scattered 




10 






Tropical 




10 






Temperate 




5 






Endemic 


• 


11 






Total . 


45 





178 ANNALS OF THE SOUTH AFRICAN MUSEUM 

Kirchenpaueria triangulata (temperate) : minimum record Agulhas Bank 111m, 

minimum east coast record 350 m. 
Sertularella leiocarpa (temperate): minimum record Tristan 183 m, minimum 

east coast record 280 m. 
Zygophylax africana (endemic) : minimum east coast record 400 m. 

For the (10) species in this eurybathic/stenothermic category the limiting 
factor is obviously temperature rather than depth. 

The remaining (23) species are truly eurybathic and eurythermic, for they 
can occur in any temperature and at any depth. As might be expected, the 
majority (13) are cosmopolitan or scattered. 

Hedgpeth (1957) suggested that all the deep-sea benthic population will 
eventually prove to be cosmopolitan, but it is clear from this analysis that 
geographically cosmopolitan species are not necessarily eurythermic. 

The data for a selected group of 21 species were examined to see whether 
the depth distribution round the coast bore any relation to changing water 
temperatures. These were the species recorded from a minimum of 10 sectors 
and from depths extending below 100 m. Only 7 of the species so tested showed 
indications of an increasing minimum depth as one passes round the coast from 
west to east and then north; only 5 showed indications of an increasing maximum 
depth (Fig. 7, Table 7). This might be interpreted as an inability in a few species 
to survive in the warmer surface and shallow waters of the east coast. It is not 
surprising that none of these species belongs to the tropical component. 

Cladocarpus is a characteristic deep-water genus, and 12 of the 13 South 
African species extend down below 100 m, 9 below 200 m, 3 below 500 m and 
7 do not occur in water shallower than 200 m. 

Table 7 

The relationship between depth and distribution. 
A. The minimum recorded depth in metres; B. The maximum recorded depth in metres. 











Sectors 










Species 
















Component 


















0-5 


6-10 


11-15 


16-20 21-24 


25-30 


31-35 




A. An fennel la quadriaurita 


— 


35 





9 


50 


100 


— ■ 


Scattered 


Halecium delicatulum . 


— 


— 





4 


27 


110 


55 


Cosmopolitan 


Halopteris tuba 


— 


— 


22 


11 


27 


70 


110 


Endemic 


Salacia articulata . 





23 





10 





18 


110 


Endemic 


Sertularella flabellum . 


— 


— 


15 


10 


27 


124 


— 


Endemic 


Sertularella megista 


— 


— 





10 


27 


49 


110 


Endemic 


Symplectoscyphus 


















arboriformis 





10 


11 


11 


49 


124 


— 


Endemic 


B. An fennel la quadriaurita 


— 


40 


79 


114 


210 


825 


— 


Scattered 


Salacia articulata . 





23 


75 


110 


91 


135 


132 


Endemic 


Sertularella flabellum . 


— 


— 


99 


200 


91 


232 


— 


Endemic 


Sertularella megista 


— 


— 


75 


200 


164 


219 


347 


Endemic 


Symplectoscyphus 


















arboriformis 


— 


20 


35 


120 


90 


420 


— 


Endemic 



THE GEOGRAPHICAL DISTRIBUTION OF SOUTHERN AFRICAN HYDROIDS 



179 



SOUTH 



EAST 




SECTORS 

Fig. 7. The distribution according to depth of three species which show tendencies towards 
greater depths in warmer waters. Actual records. 



ESTUARINE AND BRACK-WATER SPECIES 

Although many hydroids penetrate into the mouths of estuaries, especially 
on rocky substrata, only two species are truly estuarine and have not been found 
in the open sea. These are Bimeria fluminalis, which grows on mangrove roots 
in Richards Bay and St Lucia estuaries, and Hydractinia kaffraria, which is a 
commensal on the shell of Nassa kraussiana and occurs in estuaries from the 
Breede River mouth on the south coast to Durban on the east coast. 

The Limnomedusan, Ostroumovia inkermanica, is known from several 
brack-water lakes on the east coast, the hydroid from Lagoa Poelela, Mozam- 
bique, in a salinity of 8 parts per thousand, both hydroid and medusa from 
Nhlange Lake, Zululand, in 3-4 parts per thousand, and the medusa alone from 



180 



ANNALS OF THE SOUTH AFRICAN MUSEUM 



■ I 






A 


y/" 


-20 i 




• 


Lovenella chiquitita 


£ 20- 






■ 


Cladocarpus leloupi 




- 




A 


Sertularella natalensis 


1 


-25 








£*S 25- 


-30 








30- 


-35 






20 25 30 


35- 




15 
, 1 . 




35 

. , 1 . . 




-30 



-35 



■ I 




c 






S- 


-20 ) 




Synthecium 


hians 




C 20- 


-25 










^* 


-30 








4* 


30- 


- 






%*• 




■ 


-35 




k^mv+m • 






35- 


1 l 1 


15 
l 1 1 1 


20 25 
l 1 1 1 1 1 i 1 1 


30 
i i l 1 i 




35 



• I 






D 








-20*^ 






Salacia articulata 






20- 


-25 












*25- 


- 








)• 




■ 


-30 






m% 






30- 


-35 




mi 


20 25 30 
' 1 ■ ■ ■ ■ J i i i i 1 . . 






35- ; 


i i i 


15 

-J.J-.I 


■ i 


, 


35 





- L 




E 




S- 


-20*V 




Symplectoscyphus 


macrogonus 


{20- 


-25 








+r 25- 


-30 








30- 


- 




•5 j»# 




■ 


-35 








35- 


1 l l 


15 
i 1 I 


20 25 
1 1 i 1 — i — I — i — i — 1 — i 


30 

1— 1 1 1 1 — 1 L_ 


35 
1 1 l 1 




Fig. 8A-D. Distribution of six representative endemic species: A. Lovenella chiquitita 

(west coast + False Bay), Cladocarpus leloupi (south coast) and Sertularella natalensis (east 

coast). B. Corhiza scotiae (west + south coast). C. Synthecium hians (south + east coast). 

D. Salacia articulata (west + south + east coast). 

E-F. Distribution of two representative temperate species: E. Symplectoscyphus macrogonus 

(west + south coast). F. Sertularella leiocarpa (west + south + east coast), also steno- 

thermic. 



THE GEOGRAPHICAL DISTRIBUTION OF SOUTHERN AFRICAN HYDROIDS 181 



- 1 




A 


S- 


-20 ) 




Lytocarpus philippinus 


( 20- 


-25 






rf -*-^25- 


-30 






30- 


- 






- 


-35 






35- 


III 


15 

-1 1 l 


20 25 30 

i i i 1 i i i i J i i i i 1 i i 


35 

I I 1 I 1 



- I 




B 


y^ 


-20 1 




Dynamena crisioides 


C 20- 


-25 






€ *~' 25- 


■ 






i 


- 






- 


-30 






30- 


. 




Y jf% 


. 


- 






- 


- 






- 


-35 






35- 




15 
i i 1 


20 25 30 
1 1 i 1 I • . . 1 1 1 1 . I I i 


35 
i 1 1 1 1 



- 1 




c 


J* 


-20 1 




Sertularia turbinata 


\ 20- 


-25 




( 


£*** 1S- 


-30 






30- 


■ 






- 


-35 






35- 


1 l l 


15 

■ 1 i i 


20 25 30 


35 



- 1 




D 






jf- 


-20 ) 




Sertularella arbuscula 




20- 


-25 










1 25- 


-30 






mi 
m% 




30- 


- 






wm 
jfm 




- 






•\ -^**< 


!• 






-35 

' ■ ' 


15 

-J_L_L 


€*^mV9mmmm 

••• 25 


30 

1 i 1 i i i 


35 

i 1 , 


35- 

J— i — 









E 


f- 


-20 ) 






Hebella scandens 


( 20- 


-25 








-** 25- 


-30 






dm 
mm 
jfm 


30- 


-35 


15 

..i, 1 I 


# ] 


_ mmmm 
^mjnrmm m 

25 30 
j — 1 — i — i — i_i — 1 i i i i — I — i_i — 1_ 


35- 

35 
. 1 , , 



I 




F 


y^ 


-20 ^ 




Amphisbetia operculata 


f 20- 


-25 






t^ 25- 


-30 






30- 


-35 


15 
, 1 . 


cms — •• • 

* 25 30 


35- 
35 
11111 



Fig. 9A-D. Distribution of four representative tropical species from east coast : A. Lytocarpus 
philippinus (to just north of Durban). B. Dynamena crisioides (to East London). C. Sertularia 
turbinata (to Mossel Bay). D. Sertularella arbuscula (to Saldanha Bay). 
E-F. Distribution of two representative cosmopolitan species: E. Hebella scandens (St Helena 
Bay to Mozambique). F. Amphisbetia operculata (Liideritz to Natal). 



182 



ANNALS OF THE SOUTH AFRICAN MUSEUM 



Lake St Lucia North in 10,4-10,8 parts per thousand. 

CONCLUSION 

The composition of the southern African hydroid fauna is summarized in 
Table 8, whence it can be seen that there is a mixture of endemic, tropical, 
temperate and cosmopolitan species in that order of abundance. 

The presence of a large endemic population, the particular distribution of 
some of the Western Indian Ocean tropical species (p. 169), and the apparent 
proliferation of subspecies in certain genera (p. 173), all point to an active 
evolutionary centre in the South African area. 

It is clear that there has also been a heavy invasion of tropical species from 
the Indo-West-Pacific region, and these species form the major component of 
the fauna of the east coast. 

Regarding the temperate and cosmopolitan species, and considering 

(i) the low relationship of the South African temperate fauna to that of the 
antarctic and subantarctic regions on the one hand, and to that of South 
America and Australasia on the other, 

(ii) the practically certain role of ships in the introduction of at least three 
species out of 251 (1,2%) to South Africa within a period of some 300 years 
(p. 170), 



Table 8 

Summary of analysis of geographical components of South African hydroid fauna (from actual 

records, not range). 



Component 


West-plus-south 

coast: 
Sectors 1-23 
(171 species) 


East coast: 

Sectors 24-34 
(182 species) 


Whole of 
South Africa: 
Sectors 1-34 
(251 species) 




No. Percentage 


No. Percentage 


No. Percentage 


TROPICAL 

Circumtropical 
Indo-West-Pacific . 
W. Indian Ocean . 


10 5,81 
7 4,1 M9,3 
16 9,4 J 


18 9,91 

23 12,6^39,6 

31 17,0J 


20 8,01 

24 9,6 V30,7 

33 13, lj 


TEMPERATE 

Antitropical . 
Southern .... 
Atlantic .... 


10 5,81 
8 4,7^13,4 
5 2,9j 


5 2,71 
5 2,7 V6,5 
2 1,1 J 


1 1 4,41 
10 4,0^ 11,2 
7 2,8j 


ENDEMIC 

To one area only . 
To the whole of South 
Africa 


35 20,5] 

V38,0 
30 17,5 J 


17 9,31 

^25,8 
30 16,5j 


52 20,7] 

V32,7 
30 12,0 J 


UNCLASSIFIED 

Cosmopolitan 
Scattered .... 


22 12,9\ 
28 16,4 J ' 


28 15,4\ 
23 12,6/ 2 *' U 


28 11,21 

36 14,3/^' 



THE GEOGRAPHICAL DISTRIBUTION OF SOUTHERN AFRICAN HYDROIDS 183 

(iii) the fact that drifting weed and flotsam has probably performed a similar 

role throughout the ages, 
it is hardly necessary to evoke the Theory of Continental Drift to explain the 
distribution. 

REFERENCES 

Arnaud, F., Arnaud, P. M., Intes, A. & Le Loeuff, P. 1976. Transport dTnvertebres 

benthiques entre l'Afrique du Sud et Sainte Helene par les laminaires (Phaeophyceae). 

Bull. Mus. natn. Hist. nat. Paris (3) 384, Ecol. gen. 30: 49-55. 
Beurois, J. 1975. Etude ecologique et halieutique des fonds de peche et des especes d'interet 

commercial (langoustes et poissons) des iles Saint-Paul et Amsterdam (Ocean Indien). 

CNFRA 37: 1-91. 
Blanco, O. M. 1967. Contribucion al conocimiento de los hidrozoarios Argentinos. Revta 

Mus. La Plata (N.S.) Zool. 9: 243-297. 
Blanco, O. M. 1968. Nueva contribucion al conocimiento de la fauna marina Hidroide. 

Revta Mus. La Plata (N.S.) Zool. 10: 195-224. 
Blanco, O. M. 1973. Nuevos plumularidos para aguas Argentinas. Neotropica 19: 73-78. 
Blanco, O. M. 1974. Adicion a los hidrozoos Argentinos. Neotropica 20: 40-47. 
Blanco, O. M. & Bellusci de Miralles, D. A. 1972a. Nuevos aportes a los campanularidos 

de Argentina. Revta Mus. La Plata (N.S.) Zool. 11: 137-151. 
Blanco, O. M. & Bellusci de Miralles, D. A. 1 9726. Hidrozoos de lalslaPedroI. Contrnes 

Inst, ant art. argent. 145: 3-43. 
Briggs, J. C. 1974. Marine zoogeography. New York, etc.: McGraw-Hill. 
Buchanan, J. B. 1957. The hydroid fauna of the Gold Coast. Revue Zool. Bot. afr. 56: 349-372. 
Calder, D. R. 1975. Biotic census of Cape Cod Bay: hydroids. Biol. Bull. mar. biol. Lab. 

Woods Hole 149: 287-315. 
Calder, D. R. 1976. The zonation of hydroids along salinity gradients in South Carolina. 

In: Coelenterate ecology and behaviour, ed. G. O. Mackie: 165-174. New York & London: 

Plenum Press. 
Clark, A. M. & Courtman-Stock, J. 1976. The echinoderms of southern Africa. London: 

British Museum. 
Cooke, W. J. 1975. Shallow water hydroids from Enewetak Atoll, Marshall Islands. 

Micronesica 11: 85-108. 
Cornelius, P. F. S. 1975a. The hydroid species of Obelia (Coelenterata, Hydrozoa: Campanu- 

lariidae), with notes on the medusa stage. Bull. Br. Mus. nat. Hist. (Zool.) 28: 251-292. 
Cornelius, P. F. S. 19756. A revision of the species of Lafoeidae and Haleciidae (Coelenterata: 

Hydroida) recorded from Britain and nearby seas. Bull. Br. Mus. nat. Hist. (Zool.) 28: 

375-426. 
Day, J. H. 1967. A monograph on the Polychaeta of southern Africa. London: British Museum. 
Day, J. H. 1974. The ecology of Morrumbene Estuary, Mozambique. Trans. R. Soc. S. Afr. 

41:43-97. 
Ekman, S. 1953. Zoogeography of the sea. London: Sidgwick & Jackson. 
Hedgpeth, J. W. 1957. Marine biogeography. In: Treatise on marine ecology and paleo- 

ecology. I. Mem. geol. Soc. Am. 67: 359-382. 
Hulley, P. A. 1972. The origin, interrelationships and distribution of southern African 

Rajidae (Chondrichthyes, Batoidei). Ann. S. Afr. Mus. 60: 1-103. 
Mergner, H. & Wedler, E. 1977. IJber die Hydroidpolypenfauna des Roten Meeres und 

seiner Ausgange. Meteor ForschErgebn. (D) 24: 1-32. 
Michel, C. 1974. Notes on marine biology studies made in Mauritius. Bull. Maurit. Inst. 

7 (2): 1-284. 
Millar, R. H. 1971. The biology of ascidians. Adv. mar. Biol. 9: 1-100. 
Millard, N. A. H. 1952. Observations and experiments on fouling organisms in Table Bay 

harbour, South Africa. Trans. R. Soc. S. Afr. 33: 415-445. 
Millard, N. A. H. 1966. Hydroids of the Vema Seamount. Ann. S. Afr. Mus. 48: 489-496. 
Millard, N. A. H. 1975. Monograph on the Hydroida of southern Africa. Ann. S. Afr. Mus. 

68: 1-513. 



1 84 ANNALS OF THE SOUTH AFRICAN MUSEUM 

Millard, N. A. H. 1917a. Hydroids from the Kerguelen and Crozet shelves, collected by the 

cruise MD.03 of the Marion- Dufresne. Ann. S. Afr. Mus. 73: 1-47. 
Millard, N. A. H. 1911 b. The South African Museum's Meiring Naude cruises. Part 3. 

Hydroida. Ann. S. Afr. Mus. 73: 105-131. 
Millard, N. A. H. & Bouillon, J. 1975. Additional hydroids from the Seychelles. Ann. S. Afr. 

Mus. 69: 1-15. 
Penrith, M. J. 1976. Distribution of shallow water marine fishes around southern Africa. 

Cimbebasia (A) 4: 137-154. 
Penrith, M.-L. 1970. The distribution of the fishes of the family Clinidae in southern Africa. 

Ann. S. Afr. Mus. 55: 135-150. 
Penrith, M.-L. & Kensley, B. F. 1970a. The constitution of the intertidal fauna of rocky 

shores of South West Africa. Part I. — Ltideritzbucht. Cimbebasia (A) 1: 191-239. 
Penrith, M.-L. & Kensley, B. 19706. The constitution of the fauna of rocky intertidal shores 

of South West Africa. Part II. -Rocky Point. Cimbebasia (A) 1: 243-268. 
Ralph, P. M. 1961. New Zealand thecate hydroids. Part V.— The distribution of the New 

Zealand thecate hydroids. Trans. R. Soc. N. Z. (Zool.) 1: 103-111. 
Smith, J. L. B. 1949. The sea fishes of southern Africa. Cape Town: Central News Agency. 
Smith, M. M. 1970. Endemism in South African fishes. In: Oceanography in South Africa. 

Mimeographed Res. SANCOR Sympos., Durban, No. H3: 1-1 1. 
Stechow, E. 1925. Hydroiden der deutschen Tiefsee-Expedition. Wiss. Ergebn. dt. Tiefsee- 

Exped. 'Valdivia' 17: 383-546. 
Stephenson, T. A. 1947. The constitution of the intertidal fauna and flora of South Africa. 

Part III. Ann. Natal Mus. 11: 207-324. 
Stephenson, T. A. & Stephenson, A. 1972. Life between tidemarks on rocky shores. 

San Francisco: W. H. Freeman & Co. 
Thiel, H. 1962. Clavopsella quadranularia nov. spec. (Clavopsellidae nov. fam.), ein neuer 

Hydroidpolyp aus der Ostsee und seine phylogenetische Bedeutung. Z. morph. Okol. Tiere 

51 : 227-260. 
U.S. Naval Oceanographic Office. 1967. Monthly charts of mean, minimum, and maximum 

sea surface temperature of the Indian Ocean. Spec. Publ. nav. oceanogr. Off., Washington 

99: 1-48. 
Watson, J. E. 1975. Hydroids of Bruny Island, southern Tasmania. Trans. R. Soc. S. Aust. 

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to 

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3 

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3 


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3 


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THE GEOGRAPHICAL DISTRIBUTION OF SOUTHERN AFRICAN HYDROIDS 



193 























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194 



ANNALS OF THE SOUTH AFRICAN MUSEUM 



x § 

H 

PL, "a 

Si 



T3 
4> 

3 
# C 

+j 

c 
o 

cj 

X 

3 
z 

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< 



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o 2 




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H o 


H 2 


I) o 


u u 


■g« 


■gw 


w 


U4 



73-2 



■3«m 

E + 

W 



.. o 
o u 



.a J a 



C Q, 



U 



s 



§ 



^00 

yoo 



■qOv 



-a on 



"B ON 
3" 1 



B "' 

Tsos 



2^ 

sr 



% 






s 



St 



-*5 

r 



w 

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6 + 8 
to ' o 

W 



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r 



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r 



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5r- 

£2 

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a, 
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IS 

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THE GEOGRAPHICAL DISTRIBUTION OF SOUTHERN AFRICAN HYDROIDS 



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196 



ANNALS OF THE SOUTH AFRICAN MUSEUM 



•)— > 
C 

o 



Q 

Z 

UJ 

Ph 
Ph 



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to 




















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THE GEOGRAPHICAL DISTRIBUTION OF SOUTHERN AFRICAN HYDROIDS 197 





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THE GEOGRAPHICAL DISTRIBUTION OF SOUTHERN AFRICAN HYDROIDS 



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6. SYSTEMATIC papers must conform to the International code of zoological nomenclature 
(particularly Articles 22 and 51). 

Names of new taxa, combinations, synonyms, etc., when used for the first time, must be 
followed by the appropriate Latin (not English) abbreviation, e.g. gen. nov., sp. nov., comb, 
nov., syn. nov., etc. 

An author's name when cited must follow the name of the taxon without intervening 
punctuation and not be abbreviated; if the year is added, a comma must separate author's 
name and year. The author's name (and date, if cited) must be placed in parentheses if a 
species or subspecies is transferred from its original genus. The name of a subsequent user of 
a scientific name must be separated from the scientific name by a colon. 

Synonymy arrangement should be according to chronology of names, i.e. all published 
scientific names by which the species previously has been designated are listed in chronological 
order, with all references to that name following in chronological order, e.g.: 

Family Nuculanidae 

Nuculana (Lembulus) bicuspidata (Gould, 1 845) 

Figs 14-1 5A 
Nucula (Ledd) bicusphlata Gould, 1845: 37. 
Leda plicifera A. Adams, 1856: 50. 

Laeda bicuspidata Hanley, 1859: 118, pi. 228 (fig. 73). Sowerby, 1871 : pi. 2 (fig. 8a-b). 
Nucula largiilierti Philippi, 1861 : 87. 
Leda bicuspidata: Nickles, 1950: 163, fig. 301; 1955: 110. Barnard, 1964:234, figs 8-9. 

Note punctuation in the above example: 
., comma' separates author's name and year 
semicolon separates more than one reference by the same author 
full stop separates references by different authors 

figures of plates are enclosed in parentheses to distinguish them from text-figures 
dash, not comma, separates consecutive numbers 

Synonymy arrangement according to chronology of bibliographic references, whereby 
the year is placed in front of each entry, and the synonym repeated in full for each entry, is 
not acceptable. 

In describing new species, one specimen must be designated as the holotype; other speci- 
mens mentioned in the original description are to be designated paratypes; additional material 
not regarded as paratypes should be listed separately. The complete data (registration number, 
depository, description of specimen, locality, collector, date) of the holotype and paratypes 
must be recorded, e.g. : 

Holotype 

SAM-A 13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King's Beach 
Port Elizabeth (33°51'S 25°39'E), collected by A. Smith, 15 January 1973. 

Note standard form of writing South African Museum registration numbers and date. 

7. SPECIAL HOUSE RULES 
Capital initial letters 

(a) The Figures, Maps and Tables of the paper when referred to in the text 

e.g. '. . . the Figure depicting C. namacolus . . .'; \ . . in C. namacolus (Fig. 10) . 

(b) The prefixes of prefixed surnames in all languages, when used in the text, if not preceded 
by initials or full names 

e.g. Du Toit but A. L. du Toit; Von Huene but F. von Huene 

(c) Scientific names, but not their vernacular derivatives 
e.g. Therocephalia, but therocephalian 

Punctuation should be loose, omitting all not strictly necessary 
Reference to the author should be expressed in the third person 
Roman numerals should be converted to arabic, except when forming part of the title of a 

book or article, such as 

'Revision of the Crustacea. Part VIII. The Amphipoda.' 
Specific name must not stand alone, but be preceded by the generic name or its abbreviation 

to initial capital letter, provided the same generic name is used consecutively. 
Name of new genus or species is not to be included in the title : it should be included in the 

abstract, counter to Recommendation 23 of the Code, to meet the requirements of 

Biological Abstracts. 



N. A. H. MILLARD 

THE GEOGRAPHICAL DISTRIBUTION OF 
SOUTHERN AFRICAN HYDROIDS 



VOLUME 74 PART 7 



DECEMBER 1977 



ISSN 0303-2515 



OF THE SOUTH AFRICAN 

MUSEUM 



:ape town 




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1. MATERIAL should be original and not published elsewhere, in whole or in part. 

2. LAYOUT should be as follows: 

(a) Centred masthead to consist of 

Title : informative but concise, without abbreviations and not including the names of new genera or species 

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Number of illustrations (figures, enumerated maps and tables, in this order) 

(b) Abstract of not more than 200 words, intelligible to the reader without reference to the text 

(c) Table of contents giving hierarchy of headings and subheadings 

(d) Introduction 

(e) Subject-matter of the paper, divided into sections to correspond with those given in table of contents 

(f ) Summary, if paper is lengthy 

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(h) References 

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5. REFERENCES cited in text and synonymies should all be included in the list at the end of 
the paper, using the Harvard System (ibid., idem, loc. cit., op. cit. are not acceptable): 

(a) Author's name and year of publication given in text, e.g. : 

'Smith (1969) describes . . .' 

'Smith (1969: 36, fig. 16) describes ..." 

'As described (Smith 1969a, 19696; Jones 1971)' 

'As described (Haughton & Broom 1927) . . .' 

'As described (Haughton et al. 1927) . . .' 

Note: no comma separating name and year 

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within each name, with suffixes a, b, etc. to the year for more than one paper by the same 
author in that year, e.g. Smith (1969a, 19696) and not Smith (1969, 1969a). 

For books give title in italics, edition, volume number, place of publication, publisher. 

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number (only if independently paged) in parentheses, pagination (first and last pages of article). 

Examples (note capitalization and punctuation) 

Bullough, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan. 

Fischer, P.-H. 1948. Donnees sur la resistance et de le vitalite des mollusques. /. Conch., Paris 88: 100-140. 

Fischer, P.-H., Duval, M. & Raffy, A. 1933. Etudes sur les echanges respiratoires des littorines. Archs 

Zool. exp. gen. 74: 627-634. 
K.OHN, A. J. 1960a. Ecological notes on Conns (Mollusca: Gastropoda) in the Trincomalee region of Ceylon. 

Ann. Mag. nat. Hist. (13) 2: 309-320. 
Kohn, A. J. 19606. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean. 

Bull. Bingham oceanogr. Coll. 17 (4): 1-51. 
Thiele, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. In: Schultze, L. Zoologische 

und anthropologische Ergebnisse einer Forschungsreise im west lichen und zentralen Sud-Afrika 4 : 269-270. 

Jena: Fischer. Denkschr. med.-naturw. Ges. Jena 16: 269-270. 

(continued inside back cover) 



ANNALS OF THE SOUTH AFRICAN MUSEUM 
ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM 



Volume 74 Band 
December 1977 Desember 
Part 7 Deel 




PAST AND PRESENT DISTRIBUTION OF SOME 
RODENT AND INSECTIVORE SPECIES IN THE 
SOUTHERN CAPE PROVINCE, SOUTH AFRICA: 

NEW INFORMATION 

By 
D. M. AVERY 



Cape Town Kaapstad 



The ANNALS OF THE SOUTH AFRICAN MUSEUM 

are issued in parts at irregular intervals as material 
becomes available 

Obtainable from the South African Museum, P.O. Box 61, Cape Town 



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Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap 



PAST AND PRESENT DISTRIBUTION OF SOME RODENT AND 

INSECTIVORE SPECIES IN THE SOUTHERN CAPE PROVINCE, 

SOUTH AFRICA: NEW INFORMATION 

By 

D. M. Avery 

South African Museum, Cape Town 

(With 3 figures and 2 tables) 

[MS. accepted 11 August 1977] 

ABSTRACT 

Micromammalian material from archaeological sites and modern owl roosts in the Cango 
valley and Gansbaai area of the southern Cape Province provides new information concerning 
the past and present distribution of certain species of rodents and insectivores. Modern 
material from the Cango valley provides evidence of a link between widely separated popula- 
tions of several species and shows Praomys natalensis (multimammate mouse) to be co-existing 
with P. verreauxi (Verreaux's mouse). The archaeological evidence indicates that Saccostomus 
campestris (Cape pouched mouse) and Steatomys pentonyx (Cape fat mouse) arrived in the 
southern Cape only at the beginning of the Holocene about 10 000 years BP and that 
P. natalensis perhaps entered the Cango valley with the advent of farming in the area around 
200 BP. On the other hand, Tatera afra (Cape gerbil), Gerbillurus paeba (pygmy gerbil) and 
possibly Otomys unisulcatus (bush Karoo rat) disappeared from the Cango valley some time 
after about 1 600 BP. 



CONTENTS 






PAGE 


Introduction 


. 201 


Present distribution . 


205 


Past distribution 


207 


Conclusion . 


208 


Acknowledgements . 


208 


References . 


209 



INTRODUCTION 

The evidence put forward below became available during the course of a 
study of micromammalian remains from archaeological sites and comparable 
modern owl roosts in the southern Cape Province. Two areas are considered, 
the Cango valley (3 322 AC) near Oudtshoorn, and the Gansbaai area (3419CB) 
on the Walker Bay coast. For the Cango valley (Fig. 2) material was available 
from the archaeological site Boomplaas A (33°23'S 22°irE) which represents 
well in excess of 40 000 years of accumulated deposit during the Upper Pleisto- 
cene and Holocene (Deacon & Brooker 1976: 211) and from five modern owl 
roosts. Two of these, Boomplaas B and C which are adjoining rock-shelters, are 
within 100 m of Boomplaas A and in the same cliff. Two others, Nooitgedacht A 

201 
Ann. S. Afr. Mus. 74 (7) 1977: 201-209, 3 figs, 2 tables. 



202 



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Cango valley (C) and Gansbaai (G) to existing ranges. A is based on Meester & Lambrechts 

(1971) and B and D-G are based on Davis (1974). 



DISTRIBUTION OF SOME RODENT AND INSECTIVORE SPECIES 



203 




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of 200 ft contours are given. 



and B (33°22'S 22°10'E) which are also adjoining rock-shelters, are situated 
2,5 km north-west of Boomplaas, whilst the fifth site, Osgat (33°22'S 22°11'E), 
lies 1,3 km north-east of Nooitgedacht and 5 km north of Boomplaas. For the 
Gansbaai area (Fig. 3) archaeological material was available from Die Kelders 1 
(34°32'S 19°22'E), covering a period from about 80 000 BP to 45 000 BP 
(years before present) during the Middle Stone Age (MSA) and from about 
2 000 BP upwards during the Late Stone Age (LSA) (Tankard & Schweitzer 
1974: 367; Tankard 1976: 155), and from Byeneskranskop 1 (34°35'S 19°28'E) 
which was occupied from about 12 000 BP upwards (F. R. Schweitzer pers. 
comm.) and which lies some 9,5 km south-east of Die Kelders. The modern 
sample was taken from Byeneskranskop 2, a rock-shelter situated just below the 
archaeological site and set in the same cliff. Total minimum numbers of 
individuals represented in the various samples are given in Table 1 . 

It appears from Davis's (1974) distribution maps that little collecting or 
recording has been undertaken in the two areas discussed here, although 
Grindley et al. (1973) collected owl pellets from near Stanford some 15 km north 
of Gansbaai. For this reason the recent information has proved useful both for 



204 



ANNALS OF THE SOUTH AFRICAN MUSEUM 




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DISTRIBUTION OF SOME RODENT AND INSECTIVORE SPECIES 205 

consolidating known ranges and for filling in gaps in others less well known. 
Remains from archaeological contexts provide some indication of when various 
species arrived in or, in some cases, disappeared from the two areas. 

PRESENT DISTRIBUTION 

The presence of many of the species listed in Table 1 was predictable and 
in these cases the evidence merely adds another locality to an already clearly 
established range. In some cases, however, the sparseness of the existing record 
adds significance to the present information. Acomys subspinosus (Cape spiny 
mouse), Saccostomus campestris (Cape pouched mouse), Dendromus melanotis 
(grey pygmy climbing mouse), D. mesomelas (chestnut climbing mouse), 
Crocidura flavescens (red musk shrew), C. cyanea cyanea (reddish-grey musk 
shrew), Suncus varilla (lesser grey dwarf shrew) and, to some extent, Praomys 
verreauxi (Verreaux's mouse) fall into this category. The evidence confirms the 
ranges of A. subspinosus and S. campestris and also the eastward extent of the 
range of P. verreauxi as given in Davis (1974: 158, 163, 160 respectively). In the 
case of D. melanotis, D. mesomelas (Davis 1974: 166, 168) and C. flavescens 
(Meester 1963: 34) confirmation is provided for the existence of links between 
populations in the south-west and others in the south-east. For S. varilla the 
present information provides not only confirmation of similar linking but also 
constitutes a southwards extension of the known range given by Meester & 
Lambrechts (1971 : 11) (Fig. 1A). 

Further to this, and even more important, is another group of species 
which has been discovered in the Cango valley. For this group evidence of links 
between isolated populations had not previously been forthcoming. Dasymys 
incomtus (shaggy swamp rat), Mystromys albicaudatus (white-tailed rat), 
Otomys laminatus (laminate vlei rat), and O. saundersae (Saunders's vlei rat) 
comprise this group (Fig. 1D-G). It is interesting to note that Davis (1974: 
166 et seq.) predicted that the gap was more apparent than real in several cases. 
Most of these species are not frequently caught by owls (Vernon 1972: 119). 
Assuming this to represent roughly the actual proportions of animals present, 
it is likely that still more records will become available with more intensive 
work. Steatomys pentonyx (Cape fat mouse) was also unexpected in the Cango 
valley but the situation regarding Steatomys spp. is complicated by the fact that 
this genus has long been in need of revision (Davis 1962: 73). Currently the 
south-western Cape population is referred to S. pentonyx and the more northern 
material to at least two other species. It is thought most likely that the present 
material represents a considerable eastward extension of the range of S. pentonyx 
(Fig. 1C). 

Praomys natalensis (multimammate mouse) has not been thought to occur 
further west than about 24°E in the southern Cape, the more westerly area being 
occupied by P. verreauxi (Verreaux's mouse) (Fig. IB). Indeed, Davis (1974: 
160) doubted Pocock's identification of P. natalensis from Oudtshoorn, 
considering it more likely to be referable to P. verreauxi. However, material 



206 



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DISTRIBUTION OF SOME RODENT AND INSECTIVORE SPECIES 207 

collected from the Cango valley by the writer contains examples of both 
P. natalensis and P. verreauxi. Not only does this confirm Pocock's findings, but 
it appears to be the first record of the two species' co-existing. It therefore goes 
some way towards establishing the boundary between the two species, for which 
Davis (1974: 160) saw a need. This boundary now lies along a line between 
Oudtshoorn and Plettenberg Bay but it seems likely that P. natalensis will yet 
be found further west in view of its liking for agricultural land, discussed below, 
and its highly successful nature. 

Davis (1962: 62) remarked that P. natalensis is 'semi-domestic and present 
distribution is possibly dependent on having followed early human population 
movements'. It was probably early agricultural activity in particular which 
attracted it. Roberts (1951 : 472) and Shortridge (1942 : 60) note that it is common 
near cultivated land and the situation in the Cango valley seems to support this 
suggestion. At Boomplaas A microfauna has only been recovered from the 
pre-farming levels, the topmost of which is dated to 1 630 ± 50 BP (Deacon 
et al. 1976: 142). P. natalensis does not occur at all in these levels. It must, 
therefore, have arrived subsequently, possibly in the wake of European farmers 
who settled there around 200 BP. Not only does P. natalensis occur in significant 
numbers in the modern sample, but also there is a change in proportion between 
it and P. verreauxi in different parts of the valley (Table 2). At Boomplaas B 

Table 2 



Differences in the proportion of Praomys natalensis and P. verreauxi in 

See Table 1 for explanation of abbreviations. 


the Cango valley. 


BPB BPC Total NGA 


NGB Total 


Praomys natalensis .... 94 99 193 8 
Praomys verreauxi .... 13 9 22 27 


28 36 

22 49 



and C, which are situated on the hillside above the intensively farmed valley 
floor, there is a preponderance of P. natalensis, whereas at Nooitgedacht, in an 
uncultivated area, P. verreauxi tends to predominate. Whether the two species 
are in competition or whether a state of equilibrium has been reached remains 
to be seen. What is clear, however, is that P. natalensis is a recent newcomer to 
the area, apparently attracted by cultivated land. 

PAST DISTRIBUTION 

It is evident from Table 1 that the majority of species was present through- 
out the period of deposition in the caves. This presence was, however, inter- 
mittent in some cases, a fact which could well be due to a paucity of remains 
especially in the case of normally rare species such as Dasymys incomtus and 
Otomys laminatus. The former, for instance, has so far been found only from 
the lowest Holocene levels downwards at Boomplaas A but it is nevertheless 
present at Nooitgedacht; likewise at Byeneskranskop it occurs in one of the 
middle levels and then not again until the present. Several other species are not 



208 ANNALS OF THE SOUTH AFRICAN MUSEUM 

represented in the Holocene levels at Boomplaas A, but none of them is ever 
numerous and the present sample is not large. In this situation presence is 
obviously more significant than absence and further sampling may prove the 
fluctuation to be artificial. 

In other cases, however, the situation is much clearer. P. natalensis does not 
occur at all in the archaeological sample, as was mentioned above, whereas the 
opposite is true of Tatera afra (Cape gerbil), Gerbillurus paeba (pygmy gerbil) 
and Otomys unisulcatus (bush Karoo rat). The first two species occurred 
intermittently and the latter consistently throughout the Upper Pleistocene and 
Holocene levels at Boomplaas A, but are not represented in any of the modern 
samples collected at various times during the last two years from the Cango 
valley. The deserted nests of O. unisulcatus have, however, been found in the 
vicinity of Oudtshoorn (C. G. Coetzee pers. comm.) so it is possible that they 
may yet be found in the Cango valley. Two further species, Saccostomus 
campestris and Steatomys pentonyx, occur only in the Holocene and modern 
samples from the Cango valley. S. campestris does not occur in the Gansbaai 
area, but S. pentonyx has been found in the Die Kelders 1 LSA levels and at 
Byeneskranskop 1 and 2, but not in the Die Kelders 1 MSA levels. The evidence 
suggests, therefore, that these two species migrated into the southern Cape at 
the beginning of the Holocene some 10 000 or 12 000 years BP. 

CONCLUSION 

Increases in the record have been established for most of the species found. 
In some cases where previous records were widely scattered the present results 
provide significant confirmation of range. In others where the known pattern of 
distribution is markedly discontinuous the discovery of linking occurrences in 
the Cango valley shows that the disjunction is artificial. The occurrence of 
Praomys natalensis with P. verreauxi in the Cango valley is also of considerable 
interest since it establishes, at least provisionally, the boundary between the two 
species. 

Some general information has emerged concerning the movements of 
certain species. The evidence seems secure for the arrival at the beginning of the 
Holocene of Saccostomus campestris and Steatomys pentonyx and for the more 
recent arrival of Praomys natalensis, and for the recent departure of Tatera afra 
and Gerbillurus paeba, while Otomys unisulcatus has possibly retreated from the 
valley to the plain. Other species have been present, at least intermittently, from 
the Upper Pleistocene to the present. 

ACKNOWLEDGEMENTS 

Thanks are due to the following for making material available: Dr H. J. 
Deacon, University of Stellenbosch (Boomplaas A), Mr F. R. Schweitzer, 
South African Museum (Die Kelders 1 and Byeneskranskop 1), and Mr G. Avery 
and Miss K. Scott, South African Museum (Byeneskranskop 2). Mr A. Botha 
kindly pointed out owl roosts in the Cango valley. 



DISTRIBUTION OF SOME RODENT AND INSECTIVORE SPECIES 209 

REFERENCES 

Davis, D. H. S. 1962. Distribution patterns of southern African Muridae, with notes on some 

of their fossil antecedents. Ann. Cape prov. Mus. 2: 56-76. 
Davis, D. H. S. 1974. The distribution of some small southern African mammals (Mammalia: 

Insectivora, Rodentia). Ann. Transv. Mus. 29: 135-184. 
Deacon, H. J. & Brooker, M. 1976. The Holocene and Upper Pleistocene sequence in the 

southern Cape. In: Proceedings of the Southern African Society for Quaternary Research. 

Ann. S. Afr. Mus. 71: 203-214. 
Deacon, H. J., Deacon, J. & Brooker, M. 1976. Four painted stones from Boomplaas Cave, 

Oudtshoorn District. S. Afr. archaeol. Bull. 31: 141-145. 
Grindley, J., Siegfried, W. R. & Vernon, C. J. 1973. Diet of the Barn owl in the Cape 

Province. The Ostrich 44: 266-267. 
Meester, J. 1963. A systematic revision of the shrew genus Crocidura in southern Africa. 

Transv. Mus. Mem. 13: 1-127. 
Meester, J. & Lambrechts, A. von W. 1971. The southern African species of Suncus 

Ehrenberg (Mammalia: Soricidae). Ann. Transv. Mus. 27: 1-14. 
Roberts, A. 1951. The Mammals of South Africa. Johannesburg: The Trustees of 'The 

Mammals of South Africa' Book Fund. 
Shortridge, G. C. 1942. Field notes on the first and second expeditions of the Cape museums' 

mammal survey of the Cape Province: descriptions of some new subgenera and subspecies. 

Ann. S. Afr. Mus. 36: 27-100. 
Tankard, A. J. 1976. The stratigraphy of a coastal cave and its palaeoclimatic significance. 

Palaeoecol. Afr. 9: 151-159. 
Tankard, A. J. & Schweitzer, F. R. 1974. The geology of the Die Kelders Cave and environs: 

a palaeoenvironmental study. S. Afr. J. Sci. 70: 365-369. 
Vernon, C. J. 1972. An analysis of owl pellets collected in southern Africa. The Ostrich 43: 

109-124. 



6. SYSTEMATIC papers must conform to the International code of zoological nomenclature 
(particularly Articles 22 and 51). 

Names of new taxa, combinations, synonyms, etc., when used for the first time, must be 
followed by the appropriate Latin (not English) abbreviation, e.g. gen. nov., sp. nov., comb, 
nov., syn. nov., etc. 

An author's name when cited must follow the name of the taxon without intervening 
punctuation and not be abbreviated; if the year is added, a comma must separate author's 
name and year. The author's name (and date, if cited) must be placed in parentheses if a 
species or subspecies is transferred from its original genus. The name of a subsequent user of 
a scientific name must be separated from the scientific name by a colon. 

Synonymy arrangement should be according to chronology of names, i.e. all published 
scientific names by which the species previously has been designated are listed in chronological 
order, with all references to that name following in chronological order, e.g. : 

Family Nuculanidae 

Nuculana (Lembulus) bicuspidata (Gould, 1 845) 

Figs 14-1 5A 
Nucula (Lecia) bicuspidata Gould, 1845: 37. 
Leda plicifera A. Adams, 1856: 50. 

Laeda bicuspidata Hanley, 1859: 118, pi. 228 (fig. 73). Sowerby, 1871: pi. 2 (fig. 8a-b). 
Nucula largUlierti Philippi, 1861 : 87. 
Leda bicuspidata: Nickles, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9. 

Note punctuation in the above example : 

comma separates author's name and year 

semicolon separates more than one reference by the same author 

full stop separates references by different authors 

figures of plates are enclosed in parentheses to distinguish them from text-figures 

dash, not comma, separates consecutive numbers 

Synonymy arrangement according to chronology of bibliographic references, whereby 
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not acceptable. 

In describing new species, one specimen must be designated as the holotype; other speci- 
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not regarded as paratypes should be listed separately. The complete data (registration number, 
depository, description of specimen, locality, collector, date) of the holotype and paratypes 
must be recorded, e.g. : 

Holotype 

SAM-A 13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King's Beach 
Port Elizabeth (33°51'S 25°39'E), collected by A. Smith, 15 January 1973. 

Note standard form of writing South African Museum registration numbers and date. 

7. SPECIAL HOUSE RULES 

Capital initial letters 

(a) The Figures, Maps and Tables of the paper when referred to in the text 

e.g. '. . . the Figure depicting C. namacolus . . .'; '. . . in C. namacolus (Fig. 10) . . .' 

(b) The prefixes of prefixed surnames in all languages, when used in the text, if not preceded 
by initials or full names 

e.g. Du Toit but A. L. du Toit; Von Huene but F. von Huene 

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Punctuation should be loose, omitting all not strictly necessary 
Reference to the author should be expressed in the third person 
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'Revision of the Crustacea. Part VIII. The Amphipoda.' 
Specific name must not stand alone, but be preceded by the generic name or its abbreviation 

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Name of new genus or species is not to be included in the title : it should be included in the 

abstract, counter to Recommendation 23 of the Code, to meet the requirements of 

Biological Abstracts. 



D. M. AVERY 

PAST AND PRESENT DISTRIBUTION OF 

SOME RODENT AND INSECTIVORE SPECIES 

IN THE SOUTHERN CAPE PROVINCE, 

SOUTH AFRICA: NEW INFORMATION 



VOLUME 74 PART 8 JANUARY 1978 



ISSN 0303-2515 



MUS. COMP. ZOOL 
LIBRARY 

APR 2 8 19/ 



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UNIVERSITY 



ANNALS 




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5. REFERENCES cited in text and synonymies should all be included in the list at the end of 
the paper, using the Harvard System {ibid., idem, he. cit., op. cit. are not acceptable): 

(a) Author's name and year of publication given in text, e.g. : 

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Note: no comma separating name and year 

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(b) Full references at the end of the paper, arranged alphabetically by names, chronologically 
within each name, with suffixes a, b, etc. to the year for more than one paper by the same 
author in that year, e.g. Smith (1969o, 19696) and not Smith (1969, 1969a). 

For books give title in italics, edition, volume number, place of publication, publisher. 

For journal article give title of article, title of journal in italics (abbreviated according to the World list o, 
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number (only if independently paged) in parentheses, pagination (first and last pages of article). 

Examples (note capitalization and punctuation) 

Bullough, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan. 

Fischer, P.-H. 1948. Donnees sur la resistance et de le vitalite des mollusques. /. Conch., Paris 88: 100-140. 

Fischer, P.-H., Duval, M. & Raffy, A. 1933. Etudes sur les echanges respiratoires des littorines. Archs 

Zool. exp. gen. 74: 627-634. 
Kohn, A. J. 1960a. Ecological notes on Conns (Mollusca: Gastropoda) in the Trincomalee region of Ceylon. 

Ann. Mag. nat. Hist. (13) 2: 309-320. 
Kohn, A. J. 19606. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean. 

Bull. Bingham oceanogr. Coll. 17 (4): 1-51. 
Thiele, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. In: Schultze, L. Zoologische 

und anthropologische Ergebnisse einer Forschungsreise im west lichen und zentralen Sud-Afrika 4: 269-270. 

Jena: Fischer. Denkschr. med.-naturw. Ges. Jena 16: 269-270. 

(continued inside back cover) 



ANNALS OF THE SOUTH AFRICAN MUSEUM 
ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM 



Volume 74 Band 
January 1978 Januarie 
Part 8 Deel 




CRETACEOUS FAUNAS FROM ZULULAND AND 

NATAL, SOUTH AFRICA 

A FLICKIA FROM THE CENOMANIAN OF 

NORTHERN ZULULAND 

By 

WILLIAM JAMES KENNEDY 

& 
HERBERT CHRISTIAN KLINGER 



Cape Town Kaapstad 



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CRETACEOUS FAUNAS FROM ZULULAND AND NATAL, 

SOUTH AFRICA 

A FLICKIA FROM THE CENOMANIAN OF NORTHERN ZULULAND 

By 

William James Kennedy 

Geological Collections, University Museum, Oxford 

& 

Herbert Christian Klinger 

South African Museum, Cape Town 

(With 2 figures) 

[MS. accepted 11 August 1977] 

ABSTRACT 

The Lower Cenomanian Mzinene Formation of the Ndumu area of northern Zululand 
has yielded specimens of the peculiar cryptogenic dwarf ammonite Flickia Pervinquiere, 1907 
—type genus of the Family Flickiidae Pervinquiere, 1910, a group characterized by simplified, 
goniatitic sutures. The family was previously known almost exclusively from pyritic clay 
facies, and the present occurrences in a nearshore clastic sequence is unusual. The specimens 
are referred to Flickia quadrata Collignon, 1964, a species previously known only from 
Madagascar. They represent the first record of the family from Africa south of the Sahara. 



CONTENTS 



Introduction 

Systematic palaeontology 
Discussion . 
Acknowledgements . 
References . 



PAGE 

211 

212 
216 
217 

217 



INTRODUCTION 



In 1907 Pervinquiere introduced the new genus and species Flickia simplex 
for a group of diminutive (5-15 mm diameter) moderately evolute compressed, 
smooth ammonites with narrowly arched venters and extraordinarily goniatite- 
like sutures. Pervinquiere compared his specimens with the Triassic ceratites 
Nannites Mojsisovics, 1881, and Lecanites Mojsisovics, 1882, as well as con- 
sidering that they might perhaps be juveniles of Neolobites Fisher, 1882, a 
view which he rejected on the basis of fundamental differences in sutures and 
shell form. He did, however, regard Flickia as a close ally within the 'Hoplites'. 

Thirteen years later, W. S. Adkins, working on the pyritic ammonite 
faunas of similar clay facies occurring in Texas (Adkins 1920) described two 
further species, Flickia boesi Adkins (Adkins 1920: 85, pi. 1 (figs 1-3)) and 
Flickial bosquensis Adkins (Adkins 1920: 87, pi. 1 (fig. 4)). The latter, with 

211 
Ann. S. Afr. Mus. 74 (8), 1978: 211-217, 2 figs. 



212 ANNALS OF THE SOUTH AFRICAN MUSEUM 

prominent umbilical bullae, Adkins believed to represent a new genus, subse- 
quently named Adkinsia by Emile Bose, with Adkinsia adkinsi as type species 
(Bose 1928: 237, pi. 8 (figs 3-14)). Bose named three other species of Adkinsia 
in the same publication, all from the same horizon (the Del Rio Clay), all, 
however, probably being no more than intraspecific variants (A. sparcicosta 
Bose, 1928: 238, pi. 8 (figs 15-20); A. tuberculata Bose, 1928: 240, pi. 87 
(figs 21-26) and A. semiplicata Bose, 1928: 246, pi. 9 (figs 7-12)). In the same 
year Adkins (1928) introduced the Family Flickiidae (as Flickidae) to 
accommodate these two genera, without commenting on their affinities. 

With Collignon's (1928 onwards) monographic studies of the ammonite 
faunas of Madagascar, further examples of these curious genera were described, 
again as minute individuals in pyritic clay facies (Flickia pervinquierei Collignon, 
1928: 4, pi. 2 (figs 15, 15a); Flickia costellata Collignon, 1964: 23, pi. 322 
(fig. 1427) and Flickia quadrata Collignon, 1964: 23, pi. 322 (figs 1428-1429)). 

The group thus show a strong facies-linked distribution, but unlike some 
other pyritic micromorphs from clay facies which have proved to be no more 
than nuclei of large limestone individuals (e.g. many Submantelliceras Spath, 
1923 and Neopulchellia Collignon, 1929; see Casey (1965) and Kennedy & 
Hancock (1971, 1977) for discussions) they appear to be genuine micromorph 
inhabitants of offshore clay depositing environments. 

The present Zululand specimens, from lignitic nearshore siltstone sequences, 
are thus a unique occurrence which extend both facies and geographic ranges 
of these curious ammonites, as well as providing the first adult individuals to be 
described which retain all of the body chamber and the aperture. 

SYSTEMATIC PALAEONTOLOGY 

Phylum MOLLUSCA 

Class Cephalopoda Cuvier, 1797 

Subclass ammonoidea Zittel, 1884 

Order AMMONITIDA Hyatt, 1889 

Superfamily acanthocerataceae Hyatt, 1900 

Family Flickiidae Adkins, 1928 

Genus Flickia Pervinquiere, 1907 

Type species 

Flickia simplex Pervinquiere, 1907 by original designation. 

Flickia quadrata Collignon, 1964 
Figs 1-2 
Flickia quadrata Collignon, 1964: 23, pi. 322 (figs 1428-1429). 



CRETACEOUS FAUNAS FROM SOUTH AFRICA 



213 





Fig. 1. Flickia quadrata Collignon. A-H. BMNH C79976. I-K. BMNH C79841. Both 
from the Mzinene Formation, Cenomanian I, Locality 181 of Kennedy & Klinger (1975). 

AD, I, K x 2; E-H, J x 1. 



214 ANNALS OF THE SOUTH AFRICAN MUSEUM 

Holotype 

The specimen figured by Collignon (1964, pi. 322 (fig. 1428)), from the 
Lower Cenomanian of Beraketa sur Sakondry (Manera), Madagascar. 

Material 

Two specimens, BMNH C79841 and BMNH C79976, both from 
Locality 181 of Kennedy & Klinger (1975: 304), the Mzinene Formation 
exposed in hill slopes east of the road, 1 km south-east of the store at Ndumu, 
northern Zululand, 26°55 , 51 ,, S 32°18'29"E. C79841 is from Bed 5 and C79976 
from Bed 3 of this section (Kennedy & Klinger 1975, fig. 12, inset) and are of 
basal Cenomanian age (Cenomanian I). 

Dimensions 

All dimensions are in millimetres. 

D = diameter, Wb = whorl breadth, Wh = whorl height, U = umbilicus. 
Figures in parentheses are dimensions expressed as a percentage of over-all 
diameter. 

D Wb Wh Wb:Wh U 

BMNHC79976 23,8(100) 8,2(34) 7,5(32) 1,09 7,8(33) 

18,7(100) 6,7(36) 6,3(34) 1,06 6,5(35) 

Diagnosis 

A slightly depressed to slightly compressed Flickia ornamented by low, 
broad irregular prorsiradiate fold-like ribs separated by variably developed 
constrictions, the shell surface being covered in coarse striae. 

Description 

Both specimens are adult and retain complete body chambers extending for 
between half and two-thirds of the last whorl. They are preserved as internal 
moulds, with traces of the original nacreous aragonitic shell present. 

Coiling is evolute with a shallow, fairly wide umbilicus which comprises 
33-35 per cent of the diameter. The whorls expand rather slowly, and are 
slightly compressed on the inner whorls, becoming slightly depressed on the 
outer. The greatest breadth is a little below mid-flank; the umbilical wall is low, 
the shoulder rounded, inner flanks somewhat flattened and subparallel (Fig. 2A), 
outer flanks convergent with an arched, rounded venter. 

Ornament consists of rather irregular, low, broad, prorsiradiate fold-like 
ribs, which are narrow at the umbilical shoulder but widen across the flanks to 
pass across the venter as a broad, rounded swelling. The interspaces are of 
irregular width and depth, and some are accentuated into distinct constrictions. 
The shell surface (where preserved) of both ribs, interspaces and constrictions 
is covered by well-developed striae (Fig. ID, G); ribs, striae and constrictions 
all strengthen on the body chamber. The aperture (Fig. ID) is simple, entire, 



CRETACEOUS FAUNAS FROM SOUTH AFRICA 



215 



slightly flared in BMNH C79976, and preceded by a narrow rib in BMNH 
C79841 (Fig. II). 

Both specimens show the sutures (Figs IK, 2B), which are very simple and 
of basic Flickiid type, with entire lobes and saddles. There is a small median 
element to the external lobe E, a rather narrow lateral lobe L, and a shallow, 
broad U 2 . Saddle E/L is the largest, and has a flattened top; L/U 2 is smaller 
and rounded. In both specimens, the last few sutures crowd, indicating the 
specimens to be adult. 

Comparisons with other species 

Flickia simplex (Pervinquiere 1907: 214, pi. 9 (figs 2a-b, 3a-b, 4, 5a-b); 
Collignon 1928-9: 28, pi. 2 (figs 14, 14a); Collignon 1964: 23, pi. 322 
(fig. 1425) is a much more compressed species during middle, and presumably 
later growth. It is ornamented only by fine, falciform striae. 

Flickia pervinquierei (lectotype here designated is the original of Collignon 
1929, pi. 2 (figs 15, 15a); see also Collignon 1931: 76, pi. 3 (figs 21, 21a); 
Collignon 1964, pi. 322 (fig. 1426)) is also flat-sided and compressed, completely 
smooth and, according to Collignon (1929), distinctly carinate. In his original 
description of this species Collignon also referred the original of Pervinquiere's 
(1907) plate 9 (fig. 5) to Flickia pervinquierei. This specimen is, in fact, the 
holotype of Flickia simplex, being so designated by Pervinquiere in his 
explanation of the plate. 

Flickia costellata Collignon (1964: 23, pi. 322 (fig. 1427)), is also rather 
compressed but resembles Flickia quadrata in being ribbed, although these ribs 
are simple, blunt and narrow rather than broad folds with associated 
constrictions. 

All described Adkinsia species bear strong umbilical bullae, whilst Fischeuria 
Pervinquiere, 1910 are very involute, globose and smooth when young with an 





5 mm. 



Fig. 2. Flickia quadrata Collignon. A whorl section and B suture of BMNH C79841. 



216 ANNALS OF THE SOUTH AFRICAN MUSEUM 

angular umbilical shoulder. At least one species, Fischeuria pusilla Matsumoto & 
Inoma (1975: 290, pi. 42 (figs 4-5)), develops folds and constrictions on the 
body chamber. 

Representatives of neither of these genera are liable to be confused with 
Flickia. 

Occurrence 

Low Lower Cenomanian of Madagascar and the Ndumu area, Zululand. 

DISCUSSION 

Flickia was first described from north Africa, where Pervinquiere (1907) 
stated the same specimens to be of both 'Vraconien' (e.g. Upper Albian) and 
'Cenomanien' age in his description. Flickia quadrata was originally described 
from Collignon's Zone of M ant ellic eras martimpreyi (auctorum, non Coquand) 
{fide Kennedy & Hancock 1971) at his locality 478, Beraketa sur Sakondry 
(Manera), Madagascar. 

From the same locality Collignon (1964) recorded a large fauna, all as 
minute pyritic individuals, including many heteromorphs and the following 
normally coiled ammonites: Desmoceras latidorsatum (Michelin), D. lemoinei 
Collignon, Protokossmaticeras madagascariense Collignon, Forbesiceras 
largilliertianum (d'Orbigny), 'Neopulchella' gignouxi Collignon, Euhystrichoceras 
{Sakondry elld) madagascariense Collignon, Prionocycloides proratus (Coquand), 
P. besairiei Collignon, Flickia simplex Pervinquiere, F. pervinquierei Collignon, 
Neosaynoceras gazellae Pervinquiere, Mantelliceras aumalense (Coquand), 
M. mantelli(J. Sowerby), M. tuberculatum (Mantell), M. suzannae (Pervinquiere), 
M. decaryi Collignon and Acompsoceras water loti Collignon. 

This fauna is that which is well known elsewhere in Madagascar and also 
north Africa, where it is variously referred to the 'martimpreyi' Zone or 
Submantelliceras aumalense Subzone, whilst the closest North American faunas 
are approximately those of the Graysonites adkinsi Zone. 

The presence in South Africa of one of the taxa apparently restricted to this 
level in clay facies elsewhere is thus of interest, as it allows correlation with a 
facies where there are large ammonites. This is important in deciphering the 
still poorly understood relationship between pyritic micromorph faunas and 
those of limestone and sandstone facies in Europe, India and elsewhere, where 
pyritic micromorphs are either absent or unrecognized as nuclei of much bigger 
specimens (Kennedy & Hancock 1971, 1977). 

Bed 1 at locality 181 near Ndumu, concretions 1,6 m below the first 
specimen of Flickia quadrata, yielded abundant heteromorphs, especially 
Hamites and Anisoceras of the A.pseudoelegans Pictet & Campiche-^. campichei 
Spath group, indicative of the Upper Albian Stoliczkaia dispar Zone. Bed 2, 
which yielded a single F. quadrata, contained abundant Sharpeiceras, including 
specimens resembling S. /allot i (Collignon), S.florencae Spath and S. vohipalense 



CRETACEOUS FAUNAS FROM SOUTH AFRICA 2 1 7 

Collignon, rare Utaturiceras vicinale (Stoliczka), submantellicerine nuclei, and 
Mariella (Mariella) oehlerti (Pervinquiere) and varieties, and Ostlingoceras 
(Ostlingoceras) spp. Bed 3, a little higher still, yielded a further Flickia, and is 
crowded with the same M. {Mariella) and O. (Ostlingoceras). Some tens of 
metres higher in the section, a more diverse fauna with Forbesiceras, 
Mantelliceras and Hypoturrilites appears. 

These observations provide a basis for correlation of pyritic and limestone 
faunas at the base of the Cenomanian in the area, and suggest that Flickia 
comes from what is virtually the base of the Cenomanian. Sadly, the associated 
Zululand faunas are still insufficient to tell whether or not these levels are 
absent in Europe or simply represented in a different biofacies (Kennedy & 
Hancock 1977). 

ACKNOWLEDGEMENTS 

We thank the following colleagues who have advised and helped in many 
ways. Dr M. K. Howarth and Mr D. Phillips (British Museum, Natural History), 
Mr C. W. Wright (Oxford), Dr M. R. Cooper (Oxford), Dr J. M. Hancock 
(London), General M. Collignon (Moirans, Isere), Dr C. Duerden and Professor 
K. Young (Austin, Texas), and the staff of the Geological Collections, Oxford 
University Museum. 

One of us (W.J.K.) acknowledges with thanks financial support from the 
Royal Society, N.E.R.C. and the Trustees of the Sir Henry Strakosh Bequest. 

REFERENCES 

Adkins, W. S: 1920. The Weno and Pawpaw formations of the Texas Comanchean. Bull. Univ. 

Tex. econ. Geol. Tech. 1856: 1-172. 
Adkins, W. S. 1928. Handbook of Texas Cretaceous fossils. Univ. Texas Bull. 2838: 1-303. 
Bose, E. 1928. Cretaceous ammonites from Texas and northern Mexico. Bull. Univ. Tex. econ. 

Geol. Tech. 2748: 143-312. 
Casey, R. 1965. A monograph of the Ammonoidea of the Lower Greensand, part VI. 

Palaeontogr. Soc. (Monogr.): 399-546. 
Collignon, M. 1928-1929. Les cephalopodes du Cenomanien pyriteux de Diego — Suarez, 

Paleontologie de Madagascar. Annls. Paleont. 17 (1928): 136-160 (1-24); 18 (1929): 

1-56 (25-29). 
Collignon, M. 1931. La faune du Cenomanien a fossiles pyriteux du Nord de Madagascar. 

Annls. Paleont. 20: 43-104 (1-64). 
Collignon, M. 1964. Atlas des fossiles caracteristiques de Madagascar (Ammonites). XI 

(Cenomanien). Tananarive: Service Geologique. 
Kennedy, W. J. & Hancock, J. M. 1971 . Mantelliceras saxbii (Sharpe) and the horizon of the 

Martimpreyi Zone in the Cenomanian of England. Palaeontology 14: 437-454. 
Kennedy, W. J. & Hancock, J. M. 1977. Towards a correlation of the Cenomanian sequences 

of Japan and western Europe. Trans. Proc. palaeont. Soc. Japan Spec. Pap. 21: 127-141. 
Kennedy, W. J. & Klinger, H. C. 1975. Cretaceous faunas from Zululand and Natal, South 

Africa. Introduction, Stratigraphy. Bull. Br. Mus. nat. Hist. (Geol.) 25: 263-315. 
Matsumoto, T. & Inoma, A. 1975. Mid Cretaceous ammonites from the Shumarinai- 

Soeushinai Area, Hokkaido. Mem. Fac. Sci. Kyushu Univ., Ser. D, Geol. 23: 263-293. 
Pervinquiere, L. 1907. Etudes de paleontologie Tunisienne, 1. Cephalopodes des terrains 

secondaires. Mem. Carte geol. Tunisie, 438 pp. 
Wright, C. W. 1957. In: Arkell, W. J. et. al. Treatise on Invertebrate Palaeontology L. 

Mollusca (4). Geological Society of America and University of Kansas Press. 



6. SYSTEMATIC papers must conform to the International code of zoological nomenclature 
(particularly Articles 22 and 51). 

Names of new taxa, combinations, synonyms, etc., when used for the first time, must be 
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An author's name when cited must follow the name of the taxon without intervening 
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Synonymy arrangement should be according to chronology of names, i.e. all published 
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order, with all references to that name following in chronological order, e.g.: 

Family Nuculanidae 

Nuculana (Lembulus) bicuspidata (Gould, 1845) 

Figs 14-1 5A 
Nucula (Leda) bicuspidata Gould, 1845: 37. 
Leda plicifera A. Adams, 1856: 50. 

Laeda bicuspidata Hanley, 1859: 118, pi. 228 (fig. 73). Sowerby, 1871 : pi. 2 (fig. 8a-b). 
Nucula largillierti Philippi, 1861 : 87. 
Leda bicuspidata: Nickles, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9. 

Note punctuation in the above example: 

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In describing new species, one specimen must be designated as the holotype; other speci- 
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not regarded as paratypes should be listed separately. The complete data (registration number, 
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must be recorded, e.g. : 

Holotype 

SAM-A 13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King's Beach 
Port Elizabeth (33°51'S 25°39'E), collected by A. Smith, 15 January 1973. 

Note standard form of writing South African Museum registration numbers and date. 

7. SPECIAL HOUSE RULES 

Capital initial letters 

(a) The Figures, Maps and Tables of the paper when referred to in the text 

e.g. \ . . the Figure depicting C. namacolus . . .'; '. . . in C. namacolus (Fig. 10) . . .' 

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Name of new genus or species is not to be included in the title : it should be included in the 

abstract, counter to Recommendation 23 of the Code, to meet the requirements of 

Biological Abstracts. 



WILLIAM JAMES KENNEDY & 
HERBERT CHRISTIAN KLINGER 

CRETACEOUS FAUNAS FROM ZULULAND 

AND NATAL, SOUTH AFRICA 
A FLICKIA FROM THE CENOMANIAN OF 

NORTHERN ZULULAND 



VOLUME 74 PART 9 



JANUARY 1978 



ISSN 0303-2515 



MUS ' COMP. ZOOL 
LIBRARY 



HARVARD 
UNIVERSITY 



OF THE SOUTH AFRICAN 

MUSEUM 



CAPE TOWN 




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(a) Author's name and year of publication given in text, e.g. : 

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number (only if independently paged) in parentheses, pagination (first and last pages of article). 

Examples (note capitalization and punctuation) 

Bullough, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan. 

Fischer, P.-H. 1948. Donnees sur la resistance et de le vitalite des mollusques. /. Conch., Paris 88: 100-140. 

Fischer, P.-H., Duval, M. & Raffy, A. 1933. Etudes sur les echanges respiratoires des littorines. Archs 

Zool. exp. gen. 74: 627-634. 
Kohn, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon. 

Ann. Mag. nat. Hist. (13) 2: 309-320. 
Kohn, A. J. 19606. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean. 

Bull. Bingham oceanogr. Coll. 17 (4): 1-51. 
Thtele, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. In: Schultze, L. Zoologische 

und anthropologische Ergebnisse einer Forschungsreise im west lichen und zentralen Sud-Afrika 4: 269-270. 

Jena: Fischer. Denkschr. med.-naturw. Ges. Jena 16: 269-270. 

(continued inside back cover) 



ANNALS OF THE SOUTH AFRICAN MUSEUM 
ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM 



Volume 74 Band 
January 1978 Januarie 
Part 9 Deel 




A NEW SPECIES OF TORTANUS 
(CRUSTACEA, COPEPODA) FROM SOUTH AFRICA 



By 
JOHN R. GRINDLEY 



Cape Town 



Kaapstad 



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A NEW SPECIES OF TORTANUS 
(CRUSTACEA, COPEPODA) FROM SOUTH AFRICA 

By 

John R. Grindley 

School of Environmental Studies, University of Cape Town 

(With 4 figures and 3 tables) 

[MS. accepted 16 August 1977] 

ABSTRACT 

A new species of Copepoda Tortanus capensis (Calanoida, Tortanidae) is described frcm 
lagoons and estuaries on the west and south coasts of South Africa. This species appears to 
be most closely related to Tortanus recticauda Giesbrecht from the Red Sea and must be 
placed in the subgenus Atortus. 



CONTENTS 






PAGE 


Tortanus capensis sp. no v. 


219 


Occurrence . 


219 


Types . 






219 


Description 






219 


Juvenile stages 






225 


Remarks 






226 


Acknowledgements 






227 


References . 






227 



Tortanus capensis sp. no v. 

(Figs 1-4) 

Occurrence 

Numerous specimens, comprising adult males and females and copepodite 
stages of both males and females from estuaries and lagoons on the west and 
south coasts of South Africa, in salinities between 34,5% and 40,0% o . 

Types 

Type specimens from Langebaan Lagoon, 100 km north of Cape Town 
have been deposited in the South African Museum, Cape Town. Holotype, 
male, SAM-A 12227, allotype, female, SAM-A 12228, 15 paratypes, male, 
female and juvenile, SAM-A 12229, with dissected appendages mounted on 
slides. Further paratypes also from Langebaan Lagoon have been deposited in 
the British Museum (Natural History) and the United States National Museum. 

Description 

Female 

Length 2,20-2,40 mm. 

Prosome (Fig. 1B-C) slender, viewed dorsally three times as long as wide, 
characteristically shaped with a cervical groove prominent in lateral view. 

219 



Ann. S. Afr. Mus. 74 (9), 1978: 219-228, 4 figs, 3 tables. 



220 ANNALS OF THE SOUTH AFRICAN MUSEUM 

Forehead shallowly sloped. Rostral projections without filaments. Eyes 
prominent and reddish. All five pedigerous segments distinct but fourth and 
fifth partly fused. Posterior angles of metasome produced but rounded. 

Urosome (Fig. 1B-C) two-segmented, last segment partly fused with the 
caudal rami. The genital segment is swollen. 

Caudal rami (Fig. 1B-C) asymmetrical, right ramus slightly wider and 
longer, about four and a half times as long as wide distally but narrowed 
proximally, furnished with fine hairs on the distal part of their medial margins. 
Five plumose and one non-plumose caudal setae on each ramus, of which the 
second inner plumose seta is twice as long as the neighbouring setae. 

Antenna 1 (Fig. 1A) of fifteen separate segments, almost reaching the end 
of the caudal rami when extended. Apparently segments 1-6, 8-12 and 24-25 
are fused, but the segmentation of the proximal part of the antenna is somewhat 
obscure. Some further fusions are apparent in immature stages in the proximal 
segments and segments 23-25 are still fused in Stage IV. Aesthetasks appear on 
a number of segments particularly on the proximal part of the antenna. Four 
long setae are present on proximal segments and on segments 14, 16, 18 and 21 
while six terminal setae arising from an expanded end give the antennae a 
characteristic appearance. The terminal setae are all plumose and the four long 
ones are asymmetrically constricted near their bases. Some specimens have one 
or two small terminal non-plumose setae. 

The second antenna and mouthparts appear to be identical in the female 
and male except for the mandibular palp and are figured for the female 
(Fig. 3D-H). 

Antenna 2 (Fig. 3D) with basipod, bearing one lateral and one sub- 
terminal seta and a medial fringe of fine hairs. Exopod one-segmented, bearing 
six setae and a small sub-terminal fringe of bristles. Endopod two-segmented, 
bearing four terminal setae and a small sub-terminal seta. 

Mandible (Fig. 3E) with gnathal lobe heavily chitinized, bearing one large 
and four smaller teeth. Each of the smaller teeth has a supporting rib bearing a 
longitudinal row of short bristles on tiny transverse striations. Basipod of 
mandibular palp unarmed. Exopod and endopod each one-segmented in the 
male and bearing four and five terminal setae respectively. In the female the 
exopod and endopod are each two-segmented and bear five and six setae 
respectively. 

Maxilla (Fig. 3F) with gnathobase, bearing seven serrated and plumose 
spines, three plumose setae and three small non-plumose setae. Endopod one- 
segmented, bearing three barbed terminal setae, one of which has widely spaced 
plumules. 

Maxilla 2 (Fig. 3G) stout, with three endites, and indistinct indications of 
segmentation. Two of the three small lateral setae bear bristles. The three 
different setae on the medial endite bear a variety of bristles. The remaining six 
long terminal setae are flattened and bear lateral rows of spinules all medially 
orientated (see enlarged figure of tip). Small subsidiary setae are situated at the 



A NEW SPECIES OF TORTANUS FROM SOUTH AFRICA 



221 




Fig. 1 . Tortanus capensis sp. nov. Female. 

A. Antenna 1. B. Dorsal view. C. Lateral view. D. Fifth legs. E. First swimming leg. 
F. Second swimming leg. G. Third swimming leg. H. Fourth swimming leg. 



bases of the second and sixth long setae and the two barbed lateral setae. 

Maxilliped (Fig. 3H) small and three-segmented, the basal segment bearing 
two long, curvaceous setae ornamented with diverging pairs of bristles, and the 
third segment bearing a small lateral seta and three strongly curved barbed 
terminal setae. 



222 ANNALS OF THE SOUTH AFRICAN MUSEUM 

Swimming legs 1-4 (Fig 1E-H) biramous with two-segmented basipodite, 
two-segmented endopodite and three-segmented exopodite. They are similar in 
both sexes. The ornamentation of the swimming legs is represented in Table 1. 















Table 1 




























Ornamentati 


on of 


swimming 


; legs. 
















Protopod 






Endopod 








Exopod 












Leg 




1 




2 


1 




2 




1 






2 




3 






Si 


Se 


Si 


Se 


Si Se 


Si 


St 


Se 


Si 


Se 


Si 


Se 


Si 


St 


Se 


P! • 


1 











3 


3 


2 


1 


1 





1 





4 


I 


I 


P* . 


1 











3 


3 


2 


1 


1 


I 


1 


I 


5 


II 


II 


Pa • 


1 











3 


3 


2 


1 


1 


I 


1 


I 


5 


II 


II 


P 4 • 


1 











3 


3 


2 


1 


1 


I 


1 


I 


5 


II 


II 



Si, Se, St represent internal, external and terminal spines, or setae respectively. The number of 
setae is shown in arabic numerals and spines in roman numerals. 

The first leg has four less spines and one less seta than the remainder. The 
fourth leg has a fringe of fine hairs on the medial margins of the second basipod 
segment and the first exopod segment. All the setae on the swimming legs are 
plumose, and the setae on the endopods are much more slender than the 
remainder. 

The fifth pair of legs of the female (Fig. ID) are uniramous, three-segmented 
and symmetrical. The second segment bears an outer seta and the third segment 
bears a stout terminal spine and a subsidiary basal spine. The terminal spine is 
not articulated but fused to the third segment and it is finely barbed distally. 

Male 

Length 1,80-1,85 mm. 

Prosome (Fig. 2C, F) somewhat different from that of the female, with the 
head more evenly tapered but with a cervical groove like that of the female. 
Forehead shallowly sloped. Rostral filaments absent. Eyes not prominent and 
dark brown. All five pedigerous segments distinct but fourth and fifth partly 
fused. Posterior angles of metasome rounded and not produced. 

Urosome five-segmented of which the first is largest. Caudal rami (Fig. 2E) 
symmetrical and about five times as long as wide. The urosomes of several 
males were characteristically twisted to the right. 

Left antenna 1 (Fig 2A-B) fifteen-segmented and similar to that of the 
female. 

Right antenna 1 (Fig. 2D) geniculate, with fifteen separate segments, two 
of which are beyond the articulation. Segments 1-6, 19-21 and 22-25 fused. 
Prominent setae appear on proximal segments and on segments 14, 16, 18 and 
21. The development of the specialization of the geniculate antenna may be 
observed in the right antenna 1 of a stage V male copepodite (Fig. 4H). 



A NEW SPECIES OF TORTANUS FROM SOUTH AFRICA 



223 




Fig. 2. Tortanus capensis sp. nov. Male. 
A-B. Left antenna. C. Dorsal view. D. Right antenna. E. Caudal rami. F. Lateral view. 



Segments 11-12 and 23-25 are fused at that stage, but segments 18 to 22 are 
free and there are prominent spines on segments 18 and 20. It would appear 
that the spine on segment 18 forms the serrated lamella on that segment of the 
geniculate antenna of the adult male. 

Male fifth legs (Fig 3A-C) uniramous and asymmetrical with the right leg 
three-segmented and the left leg four-segmented. The first segment of the right 
leg is swollen and produced into a rounded projection at its medial distal corner. 



224 



ANNALS OF THE SOUTH AFRICAN MUSEUM 




Fig. 3. Tortanus capensis sp. no v. 

Male. A. Fifth legs. B. Right fifth leg. C. Left fifth leg. Female. D. Antenna. E. Mandible 
and palp. F. First maxillary. G. Second maxilla (end of one seta enlarged). H. Maxilliped. 



The second segment bears a large bilobed medial projection near its distal end, 
which bears two setae. The terminal segment is in the form of a sub-chelate 
claw, strongly chitinized near the tip and bearing two medial setae, and two tiny 
bristles near its tip. 

The first segment of the left leg is short. The second segment is longer and 
curved and bears a medial and an outer seta. The third segment is strongly 
curved tapers distally, and has an angular swelling proximally bearing a medial 



A NEW SPECIES OF TORTANUS FROM SOUTH AFRICA 



225 




Fig. 4. Tortanus capensis sp. nov. Juveniles. 

A. Stage IV female, dorsal view. B. Fifth legs, stage IV female. C. Fifth legs stage V female. 

D. Stage IV male, dorsal view. E. Fifth legs, stage IV male, tip enlarged. F. Stage V male, 

dorsal view. G. Fifth legs, stage V male. H. Right antenna 1, stage V male. 



seta. The terminal segment is even more strongly curved and bears a finely 
serrate terminal spine, two outer marginal spinules, two inner marginal setae, 
and a large number of tiny bristles. 
Spermatophore 300-380 ^m long. 

Juvenile stages (Fig. 4A-H) 

Copepodite stages including stages four and five of both males and females 
were obtained. The changes that occur in the development of the urosome and 
the fifth legs particularly are apparent in the figures. Lengths: Stage IV female 



226 ANNALS OF THE SOUTH AFRICAN MUSEUM 

1,45 mm, Stage V female 1,75 mm, Stage IV male 1,12 mm, Stage V 
male 1,50 mm. The ornamentation of the swimming legs of the Stage IV and 
Stage V copepodites is represented in Tables 2 and 3. 

Table 2 
Copepodite Stage IV. Ornamentation of swimming legs. 





Protopod 






Endopod 




Exopod 












1 




2 


1 


+ 


2 


] 


1 


2 


+ 


3 




Si 


Se 


Si 


Se 


Si 


St 


Se 


Si 


Se 


Si 


St 


Se 


Pi • • 


. . 1 











6 


2 


1 


1 





5 


I 


I 


p 2 . . 


. . 1 











6 


2 


1 


1 


I 


6 


II 


II 


p 3 . . 


. . 1 











5 


2 


1 


1 


I 


5 


II 


II 


p 4 . . 


. . 1 











4 


2 


1 


1 


I 


5 


II 


II 



Si, Se, St represent internal, external and terminal spines, or setae respectively. The number 
of setae is shown in arabie numerals and spines in roman numerals. 

Table 3 
Copepodite Stage V. Ornamentation of swimming legs. 





Protopod 




Endopod 








Exopod 












1 


2 


1 




2 




1 


2 




3 






Si Se 


Si Se 


Si Se 


Si 


St 


Se 


Si Se 


Si Se 


Si 


St 


Se 


Pi . 
p 2 . 
p 3 . 
p 4 . 


1 
1 
1 
1 








3 
3 
3 
3 


3 
3 
3 
3 


2 
2 
2 

2 


1 
1 

1 
1 


1 
1 I 
1 I 
1 I 


1 
1 I 
1 I 
1 I 


4 
5 
5 
5 


I 
II 
II 
II 


I 

II 
II 
II 



Si, Se, St represent internal, external and terminal spines, or setae respectively. The number 
of setae is shown in arabie numerals and spines in roman numerals. 

Remarks 

Tortanus capensis is apparently the twenty-first species of this peculiar 
genus of copepods to have been described. The genus Corynura was established 
by Brady (1883) for two species obtained by the Challenger expedition in the 
Philippine islands.' This generic name which was preoccupied was replaced by 
the name Tortanus (from tortus and anus) by Giesbrecht & Schmeil (1898). The 
following species are now included in this genus: Tortanus barbatus (Brady), 
1883 (=denticulatus Giesbrecht); Tortanus gracilis (Brady), 1883; Tortanus 
forcipatus (Giesbrecht), 1889; Tortanus recticauda Giesbrecht, 1889; Tortanus 
discaudatus (Thompson & A. Scott), 1897; Tortanus setacaudatus Williams, 
1906; Tortanus murrayi A. Scott, 1909; Tortanus brevipes A. Scott, 1909; 
Tortanus (Atortus sub. gen. nov.) tropicus Sewell, 1932; Tortanus derjugini 
Smirnov, 1935 (see Brodsky 1950); 'Tortanus sp.' Colefax, 1940 (In Dakin & 



A NEW SPECIES OF TORTANUS FROM SOUTH AFRICA 227 

Colefax 1940); Tortanus longipes Brodsky, 1948; Tortanus vermiculus Shen, 
1955; Tortanus spinicaudatus Shen & Bai, 1956; Tortanus denticulatus Shen & 
Lee, 1963; Tortanus compernis Gonzales & Bowman, 1965; Tortanus rubidus 
Tanaka, 1965; Tortanus giesbrechti Jones & Park, 1968; Tortanus scaphus 
Bowman, 1971 ; Tortanus Iophus Bowman, 1971 ; and Tortanus capensis sp. nov. 
(1977). 

Tortanus capensis is the first species of the genus to have been found 
anywhere round the coasts of Africa, south of the Red Sea. It appears to be 
most closely related to Tortanus recticauda from the Red Sea (Giesbrecht 1889) 
and Tortanus murrayi, scaphus and Iophus from the East Indies (A. Scott 1909; 
Bowman 1971), but it differs from each of them in many characters. Tortanus 
capensis may be most readily distinguished by the fifth legs of either the male or 
the female, which are quite distinct from those of any of the described species of 
Tortanus. 

Specimens of Tortanus capensis from the Swartkops estuary on the south- 
east coast of South Africa showed a few minor differences in comparison with 
the type material from Langebaan Lagoon. 

In the two-segmented urosome of the female and the character of the male 
fifth legs Tortanus capensis might be placed in Steuer's group 3 which he 
established in his 1926 revision of the genus Tortanus. This is equivalent to 
Sewell's (1932) subgenus Atortus (Bowman 1971). 

ACKNOWLEDGEMENTS 

I thank Mrs S. Grindley and Mr T. Wooldridge for assistance and the 
South African National Committee for Oceanographic Research for financial 
support. 

REFERENCES 

Bowman, T. E. 1971. Tortanus scaphus and Tortanus Iophus, new Pacific planktonic copepods, 

with notes on Tortanus murrayi (Calanoida, Tortanidae). Pacif. Sci. 25 (4): 521-528. 
Brady, G. S. 1883. Report on the Copepoda collected by H.M.S. Challenger during the years 

1873 to 1876. Challenger Rep. 1873-76 8: 1-142. 
Brodsky, K. A. 1948. T. longipes sp. nov. from the Sea of Japan. Proc. Pacif. Sci. Res. Inst. 

Fish Oceanogr. 26: 80. (In Russian.) 
Brodsky, K. A. 1950. Calanoida of the far eastern and polar seas of the U.S.S.R. Tabl. anal. 

faune U.R.S.S. Zool. Inst. Acad. Sci. 35: 1-442. (In Russian.) 
Dakin, W. J. & Colefax, A. N. 1940. The plankton of the Australian coastal waters of New 

South Wales. Publ. Univ. Sydney (Zool.) 1 (1): 1-215. 
Giesbrecht, W. 1889. Elenco dei Copepodi pelagici. Atti. R. Ac. Lincei (4) 5: 24-29. 
Giesbrecht, W. & Schmeil, O. 1898. Copepoda. 1. Gymnoplea. Tierreich 6: 1-169. 
Gonzales, J. G. & Bowman, T. E. 1965. Planktonic copepods from Bahia Fosforescente, 

Puerto Rico, and adjacent waters. Proc. U.S. nat. Mus. 117: 241-303. 
Jones, E. C. & Park, T. S. 1968. A new species of Tortanus (Calanoida) from Pago Pago 

Harbour, American Samoa. Crustaceana (Suppl. I): 152-158. 
Scott, A. 1909. The Copepoda of the Siboga expedition. Siboga Exped. Monogr. 29 (A): 1-323. 
Sewell, R. B. S. 1932. The Copepoda of Indian seas. Calanoida. Mem. Indian Mus. 10: 

400-402. 



228 ANNALS OF THE SOUTH AFRICAN MUSEUM 

Shen, C. J. 1955. On some marine crustaceans from the coastal water of Fenghsien, Kiangsu 

Province. Acta. zool. Sin. 7: 75-100. (In Chinese & English.) 
Shen, C. & Bai, S. 1956. The marine Copepoda from the spawning ground of Pneumatophorus 

japonicus (Houttuyn) off Chefoo, China. Acta. zool. Sin. 8: 177-234. 
Shen, C. J. & Lee, F. S. 1963. The estaurine copepoda of Chiekong and Kaikong Rivers, 

Kwangtung Province, China. Acta. zool. Sin. 15: 571-596. 
Steueur, A. 1926. Revision der Copepodengattung Tortanus Giesbr. Boll. Soc. Adriatica Sci. 

Nat. 29: 49-69. 
Tanaka, O. 1965. The pelagic copepods of the Izu region, Middle Japan. Systematic account 

XIII. Parapontellidae, Acartiidae and Tortanidae. Publ. Seto mar. biol. Lab. 12: 379-408. 
Thompson, I. C. & Scott, A. 1897. On the plankton collected continuously during the two 

traverses of the North Atlantic in the summer of 1897. Trans. Liverpool biol. Soc. 12: 80. 
Williams, L. W. 1906. Notes on marine Copepoda of Rhode Island. Amer. Nat. 40 (447): 

639-660. 



6. SYSTEMATIC papers must conform to the International code of zoological nomenclature 
(particularly Articles 22 and 51). 

Names of new taxa, combinations, synonyms, etc., when used for the first time, must be 
followed by the appropriate Latin (not English) abbreviation, e.g. gen. nov., sp. nov., comb, 
nov., syn. nov., etc. 

An author's name when cited must follow the name of the taxon without intervening 
punctuation and not be abbreviated; if the year is added, a comma must separate author's 
name and year. The author's name (and date, if cited) must be placed in parentheses if a 
species or subspecies is transferred from its original genus. The name of a subsequent user of 
a scientific name must be separated from the scientific name by a colon. 

Synonymy arrangement should be according to chronology of names, i.e. all published 
scientific names by which the species previously has been designated are listed in chronological 
order, with all references to that name following in chronological order, e.g.: 

Family Nuculanidae 

Nuculana (Lembulus) bicuspidata (Gould, 1 845) 

Figs 14-1 5A 
Nucula (Lcda) bicuspidata Gould, 1845: 37. 
Leda plicifera A. Adams, 1856: 50. 

Laeda bicuspidata Hanley, 1859: 118, pi. 228 (fig. 73). Sowerby, 1871 : pi. 2 (fig. 8a-b). 
Nucula largillierti Philippi, 1861 : 87. 
Leda bicuspidata: Nickles, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9. 

Note punctuation in the above example: 

comma separates author's name and year > 

semicolon separates more than one reference by the same author 

full stop separates references by different authors 

figures of plates are enclosed in parentheses to distinguish them from text-figures 

dash, not comma, separates consecutive numbers 

Synonymy arrangement according to chronology of bibliographic references, whereby 
the year is placed in front of each entry, and the synonym repeated in full for each entry, is 
not acceptable. 

In describing new species, one specimen must be designated as the holotype; other speci- 
mens mentioned in the original description are to be designated paratypes; additional material 
not regarded as paratypes should be listed separately. The complete data (registration number, 
depository, description of specimen, locality, collector, date) of the holotype and paratypes 
must be recorded, e.g. : 

Holotype 

SAM-A13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King's Beach 
Port Elizabeth (33°51'S 25°39'E), collected by A. Smith, 15 January 1973. 

Note standard form of writing South African Museum registration numbers and date. 

7. SPECIAL HOUSE RULES 

Capital initial letters 

(a) The Figures, Maps and Tables of the paper when referred to in the text 

e.g. '. . . the Figure depicting C. namacolus . . .'; '. . . in C. namacolus (Fig. 10) . . .' 

(b) The prefixes of prefixed surnames in all languages, when used in the text, if not preceded 
by initials or full names 

e.g. Du Toit but A. L. du Toit; Von Huene but F. von Huene 

(c) Scientific names, but not their vernacular derivatives 
e.g. Therocephalia, but therocephalian 

Punctuation should be loose, omitting all not strictly necessary 
Reference to the author should be expressed in the third person 
Roman numerals should be converted to arabic, except when forming part of the title of a 

book or article, such as 

'Revision of the Crustacea. Part VIII. The Amphipoda.' 
Specific name must not stand alone, but be preceded by the generic name or its abbreviation 

to initial capital letter, provided the same generic name is used consecutively. 
Name of new genus or species is not to be included in the title: it should be included in the 

abstract, counter to Recommendation 23 of the Code, to meet the requirements of 

Biological Abstracts. 



JOHN R. GRINDLEY 

A NEW SPECIES OF TORTANUS 
(CRUSTACEA, COPEPODA) FROM SOUTH AFRICA 



VOLUME 74 PART 10 



JANUARY 1978 



MUS. COMP. ZOOL 
LIBRARY 

APR 2 8 1978 

HARVAkJ 
UNIVERSITY 



ISSN 0303-2515 



ANNALS 




CAPE TOWN 



INSTRUCTIONS TO AUTHORS 

1. MATERIAL should be original and not published elsewhere, in whole or in part. 

2. LAYOUT should be as follows: 

(a) Centred masthead to consist of 

Title: informative but concise, without abbreviations and not including the names of new genera or species 

Author's(s') name(s) 

Address(es) of author(s) (institution where work was carried out) 

Number of illustrations (figures, enumerated maps and tables, in this order) 

(b) Abstract of not more than 200 words, intelligible to the reader without reference to the text 

(c) Table of contents giving hierarchy of headings and subheadings 

(d) Introduction 

(e) Subject-matter of the paper, divided into sections to correspond with those given in table of contents 

(f ) Summary, if paper is lengthy 

(g) Acknowledgements 
(h) References 

(i) Abbreviations, where these are numerous 

3. MANUSCRIPT, to be submitted in triplicate, should be typewritten and neat, double spaced 
with 2,5 cm margins all round. First lines of paragraphs should be indented. Tables and a list of 
legends for illustrations should be typed separately, their positions indicated in the text. All 
pages should be numbered consecutively. 

Major headings of the paper are centred capitals; first subheadings are shouldered small 
capitals; second subheadings are shouldered italics; third subheadings are indented, shouldered 
italics. Further subdivisions should be avoided, as also enumeration (never roman numerals) 
of headings and abbreviations. 

Footnotes should be avoided unless they are short and essential. 

Only generic and specific names should be underlined to indicate italics; all other marking 
up should be left to editor and publisher. 

4. ILLUSTRATIONS should be reducible to a size not exceeding 12 x 18 cm (19 cm including 
legend); the reduction or enlargement required should be indicated; originals larger than 
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size. A metric scale should appear with all illustrations, otherwise magnification or reduction 
should be given in the legend; if the latter, then the final reduction or enlargement should be 
taken into consideration. 

All illustrations, whether line drawings or photographs, should be termed figures (plates 
are not printed; half-tones will appear in their proper place in the text) and numbered in a 
single series. Items of composite figures should be designated by capital letters; lettering of 
figures is not set in type and should be in lower-case letters. 

The number of the figure should be lightly marked in pencil on the back of each illustration. 

5. REFERENCES cited in text and synonymies should all be included in the list at the end of 
the paper, using the Harvard System {ibid., idem, he. cit., op. cit. are not acceptable): 

(a) Author's name and year of publication given in text, e.g. : 

'Smith (1969) describes . . .* 

'Smith (1969: 36, fig. 16) describes . . .' 

'As described (Smith 1969a, 19696; Jones 1971)' 

'As described (Haughton & Broom 1927) . . .' 

'As described (Haughton et al. 1927) . . .' 

Note: no comma separating name and year 

Dagination indicated by colon, not p. 

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et al. in text for more than two joint authors, but names of all authors given in list of references. 

(b) Full references at the end of the paper, arranged alphabetically by names, chronologically 
within each name, with suffixes a, h, etc. to the year for more than one paper by the same 
author in that year, e.g. Smith (1969a, 19696) and not Smith (1969, 1969a). 

For books give title in italics, edition, volume number, place of publication, publisher. 

For journal article give title of article, title of journal in italics (abbreviated according to the World list o y 
scientific periodicals. 4th ed. London: Butterworths, 1963), series in parentheses, volume number, part 
number (only if independently paged) in parentheses, pagination (first and last pages of article). 

Examples (note capitalization and punctuation) 

Bullough, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan. 

Fischer, P.-H. 1948. Donnees sur la resistance et de le vitalite des mollusques. /. Conch., Paris 88: 100-140. 

Fischer, P.-H., Duval, M. & Raffy, A. 1933. Etudes sur les echanges respiratoires des littorines. Archs 

Zool. exp. gen. 74: 627-634. 
Kohn, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon. 

Ann. Mag. nat. Hist. (13) 2: 309-320. 
Kohn, A. J. 19606. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean. 

Bull. Bingham oceanogr. Coll. 17 (4): 1—51. 
Thiele, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. In: Schultze, L. Zoologische 

und anthropologische Ergebnisse einer Forschungsreise im west lichen und zentralen Sud-Afrika 4: 269-270. 

Jena: Fischer. Denkschr. med.-naturw. Ges. Jena 16: 269-270. 

(continued inside back cover) 



ANNALS OF THE SOUTH AFRICAN MUSEUM 
ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM 



Volume 74 Band 
January 1978 Januarie 
Part 10 Deel 




NOTES ON THE SCORPION FAUNA OF THE CAPE 

PART 3 

SOME OBSERVATIONS ON THE DISTRIBUTION 

AND BIOLOGY OF SCORPIONS ON 

TABLE MOUNTAIN 



By 



E. B. EASTWOOD 



Cape Town Kaapstad 



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NOTES ON THE SCORPION FAUNA OF THE CAPE 

PART 3 

SOME OBSERVATIONS ON THE DISTRIBUTION AND BIOLOGY OF 
SCORPIONS ON TABLE MOUNTAIN 

By 

E. B. EASTWOOD 

South African Museum, Cape Town 

(With 12 figures and 4 tables) 

[MS. accepted 18 August 1977] 

ABSTRACT 

A survey of the scorpion fauna of Table Mountain on the Cape Peninsula was carried 
out. Four of the six species recorded from this area were found. Distribution and habitat 
preferences are discussed with reference to topography and vegetation. Some observations on 
behaviour in the field and laboratory are recorded. A regional key is given. 

CONTENTS 

PAGE 

Introduction 229 

Description of the study area 230 

Materials and methods 230 

Distribution and habitat 232 

Scorpion shelters 236 

Coexistence 240 

Scorpion predators 241 

Some observations on behaviour .... 241 

Key to the scorpions of Table Mountain . . 244 

Conclusions 246 

Acknowledgements 247 

References 247 

INTRODUCTION 

Six species in two families of scorpions have been recorded from Table 
Mountain. Opisthophthalmus capensis (Herbst) was recorded by Pocock (1896), 
Purcell (1899) and Lawrence (1955); Uroplectes insignis Pocock by Pocock 
(1890, 1896), Kraeplin (1894, 1899), Purcell (1901) and Hewitt (1918); U. lineatus 
(Koch) was reported by Purcell (1901) and Kraeplin (1908); U. variegatus (Koch) 
by Pocock (1896) and Hewitt (1918), and Parabuthus capensis (Ehrenberg) by 
Hewitt (1918) and Eastwood (19776). A juvenile specimen from Signal Hill was 
identified as U. vittatus (Thorell) by Penther (1900) but it is very likely that this 
record pertains to U. lineatus. The work on Cape scorpions by early taxonomists 
was confined to morphological and geographical considerations. With later 
descriptions of new forms, a reappraisal of scorpion taxonomy has become very 
complex without the relevant background on ecology and behaviour. 

A survey was carried out by regular collecting over a period of one year. 

229 
Ann. S. Afr. Mus. 74 (10), 1978: 229-248, 12 figs, 4 tables. 



230 ANNALS OF THE SOUTH AFRICAN MUSEUM 

To complement field observations, live scorpions were kept in the laboratory. 
Reports are given on diversity and distribution, coexistence, habitats and 
shelters, and where possible notes are given on behaviour. A regional key was 
constructed for field or laboratory use. 

DESCRIPTION OF THE STUDY AREA 

Location 

Table Mountain forms the northern end of the mountainous ridge of the 
Cape Peninsula and lies approximately 33°57'S 18°25'E. 

Topography, geology and soils 

Table Mountain forms a plateau of Table Mountain Sandstone, which is 
surrounded by cliffs giving way to gentle slopes. The southern end is deeply 
dissected by the Disa stream forming the Orangekloof subarea. The sandstone 
overlies a layer of Malmesbury shale which in turn rests on Cape Granite. The 
last two formations give rise to the gentle slopes (Moll & Campbell 1976), the 
areas with which this study is primarily concerned. Soil types are dependent 
on the parent material, and those derived from the sandstone are usually 
acidic sands, the granite and shale derived soils contain more clay and are less 
acidic. The lower slopes are covered with sandstone debris which give rise to 
soils of mixed origin (Moll & Campbell 1976). 
Climate 

Table Mountain lies in the Winter Rainfall Region characterized by hot, 
dry summers and cool, wet winters. The topography plays an important part 
in the climate. Rainfall for the different subareas are as follows: 

Summit and eastern slope 1 420-1 780 mm 

Western, northern slopes and Signal Hill . . . . 889 mm 

Southern subarea 1 227 mm 

(from Moll & Campbell 1976) 

Temperature differences over the whole area are not as pronounced. 

Vegetation 

The most extensive plant community is macchia or fynbos, consisting of 
Proteaceae, Ericaceae, Restionaceae, Geraneaceae and other families. There 
are also small areas of natural forest communities as well as extensively 
cultivated alien forest vegetation consisting of Pinus spp., Eucalyptus spp., 
and Acacia spp. mainly on the eastern slopes and lower northern slopes. 
There are also areas of Hakea sp. on the western slopes. The classification 
of plant communities is given in Figure 1. 

MATERIALS AND METHODS 

Six study subareas were defined : 

1. North-facing slopes 4. East-facing slopes 

2. West-facing slopes 5. Plateau subarea 

3. Southern subarea (Orangekloof) 6. Signal Hill subarea 



NOTES ON THE SCORPION FAUNA OF THE CAPE 



23 




Built up area 
Dry proteoid 
Moist proteoid 
Tall proteoid 
Low shrub 
Forest scrub forest 
Short restoid 
Alien forest 



Fig. 1. Map of Table Mountain showing approximate boundaries of plant communities. 

Each subarea was divided into a number of transects, 0,5 km wide, extend- 
ing from the base of the mountain to the sandstone cliffs. The table or plateau 
was not divided up in this way. 

Collecting was carried out by moving up a transect and collecting samples 
by turning over surface objects such as rocks, logs and other debris, and search- 
ing under the loose bark of dead trees. Records were kept of altitude, soil 
types and vegetation. 



232 ANNALS OF THE SOUTH AFRICAN MUSEUM 

Several areas were isolated for the more detailed studies of demes or popula- 
tion aggregates of Opisthophthalmus and Uroplectes. In these areas burrow 
morphology, run morphology, feeding and coexistence of species were studied. 
In order to observe the feeding and burrowing behaviour of O. capensis, studies 
were carried out in the field where possible. In addition a number of specimens 
were kept in specially prepared boxes in the laboratory to complement field 
observations of feeding and burrowing. The morphology of burrows was studied 
by excavation and measurement. The numbers of observed specimens in relation 
to vegetation types is given in Table 1. 

Table 1 

Comparison of relative numbers of scorpions observed in each of the six study subareas and 

seven vegetation types. 

O. capensis U. lineatus U. insignis U. variegatus 

Study subarea Vegetation type 

c? 9 c? ? <? ? c? ? 

North- and west- Dry proteoid 65 27 27 31 5 4 

facing slopes, and 

Signal Hill . . . Low scrub 19 8 21 28 

South-facing 

slopes and 

Plateau subareas . Tall proteoid 12 17 

Plateau subarea . Short restoid 3 

East-facing Moist proteoid 2 3 

slopes subarea . . 

Forest scrub- 
forest 9 2 17 28 
North- and east- 
facing subareas . Alien forest 1 

Total . . 119 156 45 9 

DISTRIBUTION AND HABITAT 

One scorpionid species, viz. Opisthophthalmus capensis, was found on Table 
Mountain, while other recorded species were buthids. O. capensis has also 
been found on the lower-lying Cape Flats and recorded randomly in the Cape 
Peninsula. This and allied species extend into Namaqualand (Eastwood 1911 a). 
Another species, O. macer Thorell, extends from False Bay eastwards to Port 
Elizabeth (Lawrence 1955). Parabuthus capensis (Ehrenberg) extends from Clan- 
william in the north to Cape Town, and eastwards to Tulbagh (Eastwood 1911b). 
Another widely distributed species, P. granulatus (Hemprich & Ehrenberg), is 
found on the Cape Flats to the north and has recently been collected in the 
suburban areas north of the city. Uroplectes lineatus is reported by Lawrence 
(1955) to extend from Cape Town eastwards to Mossel Bay, and northwards 
as far as Tulbagh and Worcester. U. insignis is confined to Table Mountain. 
U. variegatus, another local species, is recorded from the Cape Peninsula and 
its immediate neighbourhood, and Langebaan in the north (Lawrence 1955). 

In the present survey of the Table Mountain study area, O. capensis was 
located all along the west-facing subarea, the Signal Hill subarea and the 



NOTES ON THE SCORPION FAUNA OF THE CAPE 



233 



N 



^ 
V 



o 
o 



o 




*DOj 



O, 



o 



o 



o 




• U. I meatus 

□ U. insignis 

■ U. variegatus 

O 0. capensis 

o 2 

km 



Fig. 2. Map of Table Mountain showing distribution of scorpions collected during the survey. 

western extremity of the north-facing subarea. There was a tendency for denies 
to be found in fairly open areas of the dry proteoid communities, i.e. below the 
400 m contour, where the slopes were not too steep and ground cover plentiful, 
but they also occurred in the low scrub communities where the ground cover 
and substratum were suitable (Fig. 3). 

U. lineatus is the most widely distributed scorpion on Table Mountain 
(Fig. 2). As well as occupying widely differing habitats it was found in all the 



234 



ANNALS OF THE SOUTH AFRICAN MUSEUM 




Fig. 3. Typical habitat of Opisthophthalmus capensis, Uroplectes lineatus and U. variegatus. 
In the foreground is the dry proteoid plant community and further back the low scrub community. 



subareas under study, and at varying altitudes where ground cover was adequate. 
The demes were not as obvious as those of O. capensis and in many cases only a 
few specimens could be found in a large area. This could be accounted for by 
the immovability of much of the ground cover, or thick scrub and grass, making 
collection difficult. Thus U. lineatus was found in the dry proteoid, low scrub, 
tall proteoid, moist proteoid, short restoid and forest scrub-forest plant com- 
munities. This species appeared to be most abundant in the dry proteoid 
communities on the north- and west-facing slopes. Only one specimen was 
found in alien forest. It will adapt to many situations as shown by the range of 
habitats it occupies, and is often reported to be found in the homes of residents 
near Table Mountain and even at the 600 and 700 m contours where access 
for the collector is allowed only by traverses along rocky ledges. 

Specimens of U. variegatus were found in the same areas as occupied by 
O. capensis in the dry proteoid communities (Fig. 2). 

U. insignis is taxonomically very near U. lineatus. Purcell (1902) regarded 
it as a distinct species. He recorded specimens from the Constantia and Newlands 
slopes of Table Mountain (east-facing subarea). In the present survey two 
substantial demes were located (Fig. 2). These were situated in the forest and 
scrub forest communities which dominate the east-facing subarea. Specimens 
were collected from under stones as well as from under the dried bark of Pinus 
pinaster and P. radiata stumps in areas which are being cleared of alien vege- 
tation (Fig. 4). 



NOTES ON THE SCORPION FAUNA OF THE CAPE 



235 




Fig. 4. Typical habitat of Uroplectes insignis and U. lineatus in the forest scrub-forest 
community of the east-facing study subarea. 




Fig. 5. Run and burrow of Opisthophthalmus capensis. The antechamber is absent in this 

particular run. 



236 



ANNALS OF THE SOUTH AFRICAN MUSEUM 



No specimens of Par abut hus capensis were found in the present survey. 
There are several specimens in the collection of the South African Museum 
collected from Devil's Peak (north-facing subarea) which lies in the dry proteoid 
community. 



SCORPION SHELTERS 

Opisthophthalmus capensis was always found to construct its burrow beneath 
some surface object. The excavation leading from the opening under the stone 
or log to the burrow entrance is the run. The structure of the run is dependent 
on the size and shape of the ground cover and nature of the substratum (Fig. 5). 
Thus the run varies in length and sometimes has two or even three openings. 
Just before the burrow entrance proper there is usually a bulge or separate 
section which could be called an 'antechamber' (Fig. 6B). Often food remains 
are found in this area and its function appears to be a feeding area, and possibly 
affords freer movement for the scorpion to turn or employ its sting. Excavation 
of some fifty burrows sometimes revealed food remains in the terminal chamber 
of the burrow itself. The burrow is usually a simple tunnel having a charac- 




run entrance 



burrow entrance 



terminal chamber 



B 




direction of slope 



burrow 



Fig. 6. A. Cross-section. B. Plan of typical run and burrow of Opisthophthalmus capensis. 



NOTES ON THE SCORPION FAUNA OF THE CAPE 



237 



teristically oval entrance penetrating the ground at about 30° from the hori- 
zontal (Fig. 6A). It often turns to right or left, if there are obstacles in the way, 
and ends in a very slightly enlarged terminal chamber. Variation is due to the 
nature of the substratum, size or sex of the inhabitant. Males construct shallower 
burrows or none at all, living only in a run. Only one specimen occupies each 
burrow. Ground cover, usually sandstone debris lying on soils of mixed origin, 
often sandy loam, under which burrows were found varied from 15 to 60 cm in 
diameter. If the stones lay on a slope the run entrances always faced downhill, 
ensuring dry runs during rainy weather. Table 2 gives the measurements of 
fifty burrows. 

Table 2 
Measurements in centimetres of fifty burrows of Opisthophthalmus capensis (36 <$<$, 14 99) 



Run length S 

9 • • 
Run width <$ 

9 • • 
Burrow entrance height $ 

9 
Burrow entrance width <$ 

9 
Burrow length <£ • 

9 . • 
Vertical depth of burrows $ 

9 



Min. 


Max. 


Mean 


9,0 


27,0 


17,2 


10,0 


22,0 


15,3 


2,6 


4,0 


3,2 


2,5 


3,4 


3,1 


1,8 


3,1 


2,9 


2,0 


3,6 


3,1 


2,1 


3,2 


2,9 


2,2 


3,0 


2,8 


0,0 


20,0 


10,6 


10,0 


21,0 


19,8 


3,5 


6,5 


5,2 


6,0 


13,5 


11,1 



Uroplectes lineatus occupies a wide range of shelters which may be classified 
as follows : . 

1. Beneath suitable stones— no evidence of any run construction, but where 
concavities form a natural shelter. 

2. Beneath stones where a run has been constructed. 

3. Beneath debris such as logs and dried organic matter. 

4. In rock crevices, or under stones lying on a flat rock surface. 

5. Beneath tree bark. 

A deme of U. lineatus in the south-facing subarea was found suitable for 
a study of run morphology. The structure of the run is dependent on the nature 
of both ground cover and substratum (Figs 7-8). The runs vary greatly in 
length and width (Table 3) and may have one or two entrances. 

U. lineatus in the east-facing subarea had the same shelters as U. insignis, 
namely the space between dried bark and stumps of pine trees (Fig. 9), although 
the two species were not found together under the same cover. Where sandstone 

Table 3 
Measurements in centimetres of thirty runs of Uroplectes lineatus. 

Min. Max. Mean 

Length of run ....... 3,5 16,5 11,3 

Width of run 0,8 2,2 1,4 

Ground cover size (diameter) ... 9,0 55,0 31,0 



238 



ANNALS OF THE SOUTH AFRICAN MUSEUM 



entrance 








\ 



Fig. 7. Diagrams showing variation of runs or scrapes of Uroplectes lineatus. 




Fig. 8. Run of Uroplectes lineatus. Usually the runs are not as well defined as this one. 



NOTES ON THE SCORPION FAUNA OF THE CAPE 



239 




Fig. 9. A section of bark removed from a pine stump, showing a specimen of Uroplectes 
insignis, several isopods and a longicorn beetle. 



debris formed the ground cover, only one specimen of U. lineatus or U. insignis 
was found under each stone, whereas up to nine specimens of these species 
were collected from a single stump. One specimen of U. lineatus was found in a 
shallow, wide burrow in the north-facing subarea and not covered by any 
surface object. Presumably the burrow was excavated by another animal. 

U. variegatus was found to occupy narrow runs, usually well defined with 
a small terminal chamber. Some specimens were also found in natural depres- 
sions beneath stones. Table 4 gives measurements of five runs of U. variegatus. 



Table 4 

Measurements in centimetres of five runs of Uroplectes variegatus. 

Min. Max. Mean 

Length 4,0 13,0 9,4 

Width 0,75 1,4 0,92 

Ground cover size (diameter) . . . 16,0 48,0 37,0 



Parabuthus capensis was not collected during the survey, but has been 
collected by the author from other parts of the Cape. This species often excavates 
runs which vary according to the nature of ground cover and substratum and 
usually consist of a more or less round or oval space beneath the ground cover 
with one or two entrances. 



240 ANNALS OF THE SOUTH AFRICAN MUSEUM 

COEXISTENCE 

The following pairs of species were found to coexist: Opisthophthalmus 
capensis— Uroplectes lineatus, O. capensis—U. variegatus, U. lineatus—U. 
insignis. In field studies demes or population aggregates were roughly delineated 
by markers to determine the spatial distribution of their shelters. 

Of eight demes of O. capensis, five showed a spatial overlap with demes of 
U. lineatus (Fig. 10) in studies carried out in the west-facing subarea. Specimens 
of U. variegatus were recorded in association with O. capensis in two of these 
demes where U. lineatus was absent. In one instance a specimen of U. variegatus 
had its run under the same rock as O. capensis, their respective entrances 
opening on opposite sides. 













• 




•• 










• 






• 




400 m contour 




o 


• 


• 


• 








low scrub 




o 

o o 

o 

o 


o 

8° 




• • 

• 

CP 


o° 





• 






dry proteoid 

$ U. lineatus 
O O.capesis 



Fig. 10. Diagram of population aggregates of Opisthophthalmus capensis and Uroplectes 
lineatus in the west-facing study subarea, showing spatial overlap. 

In a study of coexistence of scorpions by Williams (1970) two mechanisms 
were found to permit coexistence : the choice of different-sized prey, and habitat 
specialization. The first mechanism undoubtedly operates in the case of O. 
capensis—U. lineatus and O. capensis—U. variegatus. 

U. insignis and U. lineatus were found under the bark of pine stumps in 
the same immediate vicinity, but specimens of either species did not share the 
same shelter. The mechanism permitting coexistence in this case is not quite 
•clear. 

The record of Parabuthus capensis from Devil's Peak indicates the prob- 
ability of the coexistence of this species with U. lineatus. P. capensis from the 
Citrusdal district has been found in association with U. carinatus (Pocock) 
and O. pallidipes Koch. 



NOTES ON THE SCORPION FAUNA OF THE CAPE 241 

SCORPION PREDATORS 

Because of the nocturnal habits of scorpions, predation is not often 
reported. The author has seen a troop of baboons on the northern slopes of 
Table Mountain foraging beneath stones. Baboons (Papio ursinus (Kerr)) are 
well-known predators of scorpions. The bokmakierie (Telophorus zeylonus (Linn.)) 
and the fiscal shrike (Lanius collaris Linn.) have been reported to prey on 
Uroplectes lineatus (I. G. Taylor pers. comm.). Owls are undoubtedly the 
commonest predators of scorpions. The spotted Eagle Owl {Bubo africanus 
(Tern.)), a local species, is reported to feed on scorpions (Roberts 1940). The 
Barn Owl {Tyto alba (Scopoli)) feeds on buthids and scorpionids (Vernon 1972). 

SOME OBSERVATIONS ON BEHAVIOUR 

The behaviour exhibited in burrow construction has been discussed by 
Williams (1966) for the Neotropical species Anuroctonus phaeodactylus (Wood) 
(Vejovidae), and Newlands (1972) reported on the burrowing behaviour of 
Protophthalmus holmi Lawrence, Opisthophthalmus concinnus Newlands and 
O. flavescens Purcell, all Ethiopian Scorpionidae. 

An important adaption of Opisthophthalmus capensis is its ability to con- 
struct burrows. In the laboratory specimens made burrows by loosening tightly- 
packed soil with a biting action of the chelicerae. The loose sand was then 
scraped out backwards by legs I and II which were tucked under the body. 
Legs III and IV provided the traction while the pedipalps were extended 
anteriorly and acted as supports to prevent the body tipping forward and 
occasionally as levers which aided in the backward movement. The cauda 
was extended straight out behind the body. The soil was transported rapidly 
backward in a single movement and the soil deposited outside the entrance on a 
tumulus by a slight flick of the first two pairs of legs so that the soil sprayed out. 
Occasionally the scorpion moved its load of soil by moving backward for a few 
centimetres, stopping, and thus pulling the soil backward with each step. With 
each step there was a pause while the hind legs were moved backward again 
to repeat the process. Thus two methods were employed to transport the soil : 
the first was used in exceptionally dry soil where backward movement was 
smooth and unimpeded; the second method was employed when the soil was 
damp and therefore more difficult to move, or if the burrow was steep or filled 
with small stones, twigs and similar obstacles. Several specimens did not burrow 
but made runs under stones provided for this purpose. Initially the animal 
would remove the soil in the usual manner as well as forming a broad chamber 
beneath the stone by pushing the soil with the tail. The curved tail was laid flat 
on the surface and then extended, thus moving a good volume of soil each time. 
This tail scraping is reported as being a widespread phenomenon among 
scorpions (Newlands 1972). It is also reported as a part of burrow maintenance 
in Anuroctonus phaeodactylus (Wood) by Williams (1966). 

Field observations showed that immature specimens were adept at burrow- 



242 ANNALS OF THE SOUTH AFRICAN MUSEUM 

ing. Several second or third instar nymphs were found in burrows of 1-2 cm 
in length under stones. These burrows rarely had the characteristic oval opening 
and were often nothing more than shallow depressions. 

The tarsi of O. capensis are adopted for a pelophilous habitat, that is 
the claws are sharply curved and tarsal setae are not as dense as in O. wahlbergi 
(Thorell) for example, a psammophilous species. Caudal segment V is heavily 
sclerotized with numerous setae and well-developed keels to facilitate tail- 
scraping. Collections made in the field during the winter months (June-Sep- 
tember) indicate that these scorpions remained deep in their burrows on cold 
days, but if the sun was hot they were often found at the entrances of their 
burrows or in the runs, facing outward. This 'doorkeeping' is presumably 
maintained while the scorpion waits for prey which may seek refuge under 
stones (Alexander & Ewer 1958). The tail was curled around sideways, parallel 
to and resting on the substratum so that the telson, which was folded over the 
dorsal surface of caudal segment V, lay against the lateral side between the 
sixth and seventh mesosomal segments. From this position the metasoma could 
easily be brought into action sideways, as had been observed in the laboratory. 
This sideways movement is obviously more suitable for stinging prey while 
the scorpion is in its run. The pedipalps were arranged so that the femora were 
perpendicular to the body and the tibiae and chelae lay parallel to the body. 
The movable and immovable fingers were kept about 5 mm apart. If the scorpion 
was just inside the burrow entrance, the chelae may have protruded and were 
arranged as above. Sometimes the chelae were folded over each other with the 
dorsal and outer sections facing outwards, forming a sort of shield. This is a 
typical defensive stance for Scorpionidae (Newlands 1969). When a stone was 
lifted while a scorpion was in its run it would lift its tail so that the telson lay 
above the first caudal segment and the body was lifted ofT the substratum. If 
further aggravated it would adopt an aggressive stance so that the last two 
mesosomal segments were curved upward, the metosoma perpendicular to the 
ground except for the last caudal segment which was held horizontal with the 
telson above the last mesosomal segment or carapace. Another reaction was 
retreat, in which case the cauda was brought down backwards and the animal 
scuttled down its burrow, either backward or forward. 

The function of the doorkeeping seems to be related to feeding, where 
the scorpion waits for prey to come into the run. At no time was it observed 
that the scorpion was an active predator. However, active foraging in the 
vicinity of burrows is known for Opisthophthalmus (G. Newlands, pers. 
comm.). Feeding was observed in the field in the runs or antechambers. 

Feeding behaviour has been discussed for Neotropical scorpions by Baerg 
(1954) and Hadley & Williams (1968) (Buthidae and Vejovidae) and for the 
South African Opisthophthalmus latimanus by Alexander (1972). In the field 
O. capensis was observed feeding on Dorylinus helvolus (Formicidae) and 
Temnopteryx phalerata (Blattariae). In the laboratory a wide range of prey 
species were accepted and eaten, while others were ignored. Prey species which 



NOTES ON THE SCORPION FAUNA OF THE CAPE 243 

were accepted were Periplanita americana, Temnopteri phalerata (Blatteriae), 
Dorylinus helvolus (Formicidae), Holopternia valga (Hemiptera) as well as 
U. lineatus, isopods, Scarabaeidae, beetle larvae, moths and moth larvae. The 
following species were either killed or ignored but not eaten: Chilopoda, 
Diplopoda and Anthia spp. (Carabidae). 

During excavation of burrows food remains in the form of pellets were 
sometimes found in the terminal chambers or in the runs. The food remains 
were identified as small tenebrionid beetles and cockroaches from the indigestible 
elytras and legs. Thus feeding occurred both in the burrows and the runs. 
Feeding behaviour was elicited in the laboratory by placing the prey near the 
scorpion which usually rests in the typical doorkeeping posture. Small prey 
was firmly grasped in both chelae and the metasoma brought upright. Small 
prey was not stung and the biting action of the chelicerae began immediately. 
If the prey was lively but not very large the scorpion brought the telson into 
play from the side and usually slowly, seeking for a soft spot in which to inject 
the venom. If the prey was large and aggressive the scorpion initiated offensive 
behaviour and the telson was brought swiftly over the carapace to deliver a 
series of stings. The ingestion process lasted from 1 to 3 hours in the laboratory, 
depending on the size of the prey. The formation of pellets of food remains 
was not observed. Ingestion started immediately after capture and in heavily 
sclerotized prey the starting point was ventrally between head and thorax. The 
chelicerae moved alternately in and out while the fingers grasped at the food, 
giving rise to the typical biting action. 

Sponge-bathing activities were not observed after feeding, nor after burrow- 
ing in wet soil when the legs and chelicerae became dirty. 

In humid laboratory environments O. capensis was never seen to approach 
the dish of water in the cage and it seems likely that water balance is maintained 
by contact with a wet substratum as reported by Hadley & Williams (1968) in 
observations at night of desert scorpions. If the cage was kept very dry and 
water placed in the cage the scorpions would climb over the edge of the dish 
and dip the chelicerae and pedipalps into the water. At the same time the pedi- 
palps moved rhythmically up and down. 

Uroplectes lineatus adopted a very characteristic resting posture while 
under ground cover. The pedipalps were usually held so that they lay slightly 
anterior to or lateral to and subparallel to the carapace. The tail was curved 
over the abdomen dorsally so that the second caudal segment was horizontal 
to the body and the terminal segments curved around sideways and the telson 
lay laterally between mesosomal segments V and VI, being folded over against 
caudal segment V dorsally. In the defensive posture the tail lay behind the body 
with the telson curved forward so that it lay above the first caudal segment. 
When agitated this scorpion can strike very rapidly to the side, forward or 
backward. This species did not use the chelae to grasp an aggressor but only 
struck out with the tail. In the aggressive posture there was a slight forward 
stilting. Venom-dripping was associated with aggressive behaviour. In the 



244 ANNALS OF THE SOUTH AFRICAN MUSEUM 

laboratory it was noticed that U. lineatus was able to excavate a run, but the 
behaviour of digging was not observed. In the east-facing subarea specimens 
were found in association with Isopoda, geckos, longicorn beetles, Lagria 
villosa F. (Lagriidae), Pyrrhocoridae, Hemiptera, cockroaches {Temnopteryx 
phalerata (Sauss)) and U. insignis. In this environment both U. lineatus and 
U. insignis were observed feeding on cockroaches. Prey capture was not observed 
here or in the laboratory. 

Two pregnant female specimens of U. insignis and one of U. lineatus were 
kept in the laboratory. Parturition was observed in both species which behaved 
similarly. Parturition behaviour was as follows: the scorpion stilted on the two 
last pairs of legs prior to delivery; the first two pairs were held under the body 
so that the distal segments were parallel to the body. This posture was main- 
tained throughout parturition. The young began passing through the genital 
opening head first, one by one into the 'birth basket' formed by the first two 
pairs of legs. The young became active after a few minutes and began ascending 
the mother's back by way of her walking legs. When all the young had reached 
the back they assumed a completely random orientation, sometimes in several 
layers, which continued to the first moult. This random orientation of first 
instar young is characteristic of the Buthidae (Williams 1969) (Figs 11-12). 
With the onset of the second instar stage the litter became distinctly more 
elongate. Duration of first instar stage of U. insignis was 9-12 days with litters 
of 12 and 13 specimens, and duration of first instar of U. lineatus was 11 days 
with a litter of 8 specimens. Parturition took place from January to February. 

KEY TO THE SCORPIONS OF TABLE MOUNTAIN 

1. Chelae slender; the dorsal surface not separated by a keel (Family 
Buthidae). Found in simple runs or scrapes under stones or beneath 

tree bark . . . . . . . . . . . . . . . . . . 2 

- Chelae large, the dorsal surface separated into inner and outer sections 
by a smooth keel (Family Scorpionidae). Found in burrows, usually 
opening under stones . . . . Opisthophthalmus capensis (Herbst) 

2. Species large; stridulatory areas on dorsal surface of caudal segments 
I and II well developed. Posterior dorsal surface of caudal segment II 
raised to form a subtriangular lip Parabuthus capensis (Ehrenberg) 
Species small ; stridulatory areas on cauda absent . . . . . . 3 

3. Tergites with short lateral keels and seven black longitudinal lines 

Uroplectes variegatus (Koch) 

- Tergites without lateral keels, darkly pigmented . . . . . . 4 

4. Tubercle below aculeus distinctly laterally compressed; vesicle 
yellow . . . . . . . . . . Uroplectes lineatus (Koch) 

- Tubercle below aculeus conical and blunted; vesicle black 

Uroplectes insignis Pocock 



NOTES ON THE SCORPION FAUNA OF THE CAPE 



245 




Fig. 11. First instar of Uwplectes insignis. Note the random orientation typical of the 

Buthidae. 




Fig. 12. Second instar of Uroplectes insignis. The young are beginning to leave the mother's 

back. 



246 ANNALS OF THE SOUTH AFRICAN MUSEUM 

CONCLUSIONS 

Four scorpions species were found on Table Mountain and Signal Hill. Two 
species, Opisthophthalmus capensis and Uroplectes lineatus, were very abundant 
whereas the other two were rare (U. insignis and U. variegatus). O. capensis 
was abundant on the western north-facing subarea, the greater proportion 
occurring in the dry proteoid plant community below the 400 m contour. 
Fewer specimens occurred in the low scrub communities since these areas are 
very steep and the substratum and ground cover unstable during the rainy 
season. U. lineatus was found to be fairly abundant in all subareas studied and 
has a random distribution, occurring in varied habitats. U. variegatus and 
U. lineatus were found respectively to coexist with O. capensis, although these 
two former species were not found to coexist. The fact that all the young pro- 
duced by two females of U. insignis had the characteristics of this species 
suggests that it is a true-breeding species which coexists with U. lineatus. 
Apparently isolating mechanisms have been acquired for this example of 
sympatric association, but what these are, is not quite clear. 

The notes on the behaviour make comparison with other species possible. 
In the burrowing behaviour of O. capensis two methods are employed to 
transport the soil ; the first is similar to the behaviour shown by Protophthalmus 
holmi Lawrence (as reported by Newlands (1972)), i.e. the soil is transported 
rapidly backward in a single movement, and the second is like that of O.flaves- 
cens Purcell and O. concinnus Newlands, i.e. the soil is transported backward by 
a series of jerks. Observations of laboratory specimens of O. capensis revealed 
that the first method was used in exceptionally dry soil where backward move- 
ment was smooth and unimpeded. The second method was employed when the 
soil was damp and therefore difficult to move, or if the burrow was steep or 
filled with small stones, twigs or similar obstacles. Thus O. capensis, which is 
pelophilous, exhibits burrowing behaviour similar to psammophilous scorpions 
in certain conditions. The excavation of a run by Parabuthus capensis was 
described by Eastwood (1917b). Part of this process was similar to the behaviour 
shown by O. capensis when obstacles had to be removed, but loose sand was 
excavated in a manner characteristic to Parabuthus. The defensive behaviour 
of O. capensis is the same as the generalized behaviour for Opisthophthalmus as 
discussed by Newlands (1969). The resting posture of O. capensis prior to 
feeding differs from that of vejovid scorpions in the position of the metasoma. 
In O. capensis the metasoma is usually curved forward alongside the body 
whereas the vejovid scorpions discussed by Hadley & Williams (1968) held the 
metasoma straight out backwards. Feeding behaviour of O. capensis was 
similar to that observed for Parabuthus capensis, except that the latter was more 
reluctant to attack larger and more active prey. 

Birth activities of Neotropical Buthidae were discussed by Williams (1969) 
and this was found to be similar for U. insignis and U. lineatus. The first instar 
young of these two species showed the random orientation on the mother's 



NOTES ON THE SCORPION FAUNA OF THE CAPE 247 

back which was reported for the genus Centruroides (Williams 1969). 

Because of the position of Table Mountain, the human impact on its ecology 
is considerable. Frequent fires cause damage to the floral and faunal ecology. 
In this study it was observed that in recently burnt areas O. capensis was abun- 
dant, where U. lineatus was more abundant in adjacent unburned areas. It is 
obvious that O. capensis is able to withstand a veld fire because of its ability 
to burrow, whereas a local deme of Uroplectes can be exterminated. The lower 
slopes of the north-facing and east-facing subareas are covered in plantations 
of alien forest. In these areas there is very little insect life and consequently 
few scorpions were found. 

ACKNOWLEDGEMENTS 

I thank Dr Y. Whitehead for advice and encouragement during this project, 
and the Director of the South African Museum for providing research and 
transport facilities. For constructive comments and advice on this and other 
projects I thank Mr B. Lamoral of the Natal Museum and Mr G. Newlands 
of the South African Institute for Medical Research. 

Thanks are also due to Dr V. Whitehead, Messrs A. Byron and S. Rice 
for help with the photographic work, and the Cape Town City Engineer's 
Department and the State Department of Forestry for permits and permission 
to collect on Table Mountain. 

REFERENCES 

Alexander, A. J. 1972. Feeding behaviour in Scorpions. S. Afr. J. Sci. 68: 253-256. 
Alexander, A. J. & Ewer, D. W. 1958. Temperature adaptive behaviour in the scorpion 

Opisthophthalmus latimanus Koch. J. exp. Biol. 35: 349-359. 
Baerg, W. J. 1954. Regarding the biology of the common Jamaican scorpion. Ann. ent. 

Soc. Am. 47: 272-276. 
Eastwood, E. B. 1977a. Notes on the scorpion fauna of the Cape. Part 1. Description of 

neotypes of Opisthophthalmus capensis (Herbst) and remarks on the O. capensis and 

O. granifrons Pocock species-groups (Arachnida, Scorpionida, Scorpionidae). Ann. S. Afr. 

Mus. 72: 211-226. 
Eastwood, E. B. 1971b. Notes on the scorpion fauna of the Cape. Part 2. The Parabuthus 

capensis (Ehrenberg) species-group; remarks on taxonomy and bionomics (Arachnida, 

Scorpionida, Buthidae). Ann. S. Afr. Mus. 73: 199-214. 
Hadley, N. F. & Williams, S. C. 1968. Surface activities of some North American scorpions 

in relation to feeding. Ecology 49(4) : 726-734. 
Hewitt, J. 1918. A survey of the scorpion fauna of South Africa. Trans. R. Soc. S. Afr. 6: 

89-192. 
Kraeplin, K. 1894. Revision der Skorpione. Jb. hamb. wiss. Anst. 12: 75-96. 
Kraeplin, K. 1899. Scorpiones und Pedipalpi. Das Tierreich 8: 1-265. 
Kraeplin, K. 1908. Skorpione und Solifugen. Ergebnisse e. Forsch. Sudafrika Bd 1, Lief 2. 

Denksch. med.-naturw. Ges. Jena 13: 247-282. 
Lawrence, R. F. 1955. Solifugae, Scorpions and Pedipalpi, with checklists and keys to South 

African families, genera and species. S. Afr. anim. Life 1: 152-262. 
Moll, E. J. & Campbell, B. M. 1976. Table Mountain. A conservation and management 

report. Cape Town : University of Cape Town. 
Newlands, G. 1969. Scorpion defensive behaviour. Afr. wild Life 23: 147-153. 
Newlands, G. 1972. Notes on psammophilous scorpions and a description of a new species 

(Arachnida: Scorpionides). Ann. Transv. Mus. 27: 241-254. 



248 ANNALS OF THE SOUTH AFRICAN MUSEUM 

Penther, A. 1900. Zur kenntnis der Arachnidenfauna Siidafrikas (Scorpiones). Annln. naturh. 

Mus. Wien 15: 153-163. 
Pocock, R. I. 1890. A revision of the genera of scorpions of the family Buthidae, with descrip- 
tions of some South African species. Proc. zoo). Soc. Lond. 1890: 114-141. 
Pocock, R. I. 1896. A further revision of the species of scorpions belonging to the South 

African genera Uroplectes, Lepreus and Tityolepreus. Ann. Mag. nat. Hist. (6) 17: 377-393. 
Purcell, W. F. 1 899. On the species of Opisthophthalmus in the collection of the South African 

Museum, with descriptions of some new forms. Ann. S. Afr. Mus. 1: 131-180. 
Purcell, W. F. 1901. On some South African Arachnida belonging to the orders Scorpiones, 

Pedipalpi and Solifugae. Ann. S. Afr. Mus. 2: 137-225. 
Roberts, A. 1940. Birds of South Africa. Revised by McLachlan, G. R. & Liversidge, R. 

3rd ed. 1970. The Trustees of the John Voelcker Bird Book Fund. 
Vernon, C. J. 1972. An analysis of owl pellets collected in southern Africa. Ostrich 43: 109-124. 
Williams, S. C. 1966. Burrowing activities of the scorpion Anuroctonus phaeodactylus (Wood) 

(Scorpionida: Vejovidae). Proc. Calif. Acad. Sci. 34: 419-428. 
Williams, S. C. 1969. Birth activities of some North American scorpions. Proc. Calif. Acad. 

Sci. 37: 1-24. 
Williams, S. C. 1970. Coexistence of desert scorpions by differential habitat preference. 

Pan-Pacif Ent. 46: 254-267. 



6. SYSTEMATIC papers must conform to the International code of zoological nomenclature 
(particularly Articles 22 and 51). 

Names of new taxa, combinations, synonyms, etc., when used for the first time, must be 
followed by the appropriate Latin (not English) abbreviation, e.g. gen. nov., sp. nov., comb, 
nov., syn. nov., etc. 

An author's name when cited must follow the name of the taxon without intervening 
punctuation and not be abbreviated; if the year is added, a comma must separate author's 
name and year. The author's name (and date, if cited) must be placed in parentheses if a 
species or subspecies is transferred from its original genus. The name of a subsequent user of 
a scientific name must be separated from the scientific name by a colon. 

Synonymy arrangement should be according to chronology of names, i.e. all published 
scientific names by which the species previously has been designated are listed in chronological 
order, with all references to that name following in chronological order, e.g.: 

Family Nuculanidae 

Nuculana (Lcmbulus) bicuspidata (Gould, 1 845) 

Figs 14-15A 
Nucula (Leda) bicuspidata Gould, 1845: 37. 
Leda plicifera A. Adams, 1856: 50. 

Laeda bicuspidata Hanley, 1859: 118, pi. 228 (fig. 73). Sowerby, 1871 : pi. 2 (fig. 8a-b). 
Nucula largillierti Philippi, 1861 : 87. 
Leda bicuspidata: Nickles, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9. 

Note punctuation in the above example: 

comma separates author's name and year 

semicolon separates more than one reference by the same author 

full stop separates references by different authors 

figures of plates are enclosed in parentheses to distinguish them from text-figures 

dash, not comma, separates consecutive numbers 

Synonymy arrangement according to chronology of bibliographic references, whereby 
the year is placed in front of each entry, and the synonym repeated in full for each entry, is 
not acceptable. 

In describing new species, one specimen must be designated as the holotype; other speci- 
mens mentioned in the original description are to be designated paratypes; additional material 
not regarded as paratypes should be listed separately. The complete data (registration number, 
depository, description of specimen, locality, collector, date) of the holotype and paratypes 
must be recorded, e.g.: 

Holotype 

SAM-A 13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King's Beach 
Port Elizabeth (33°51'S 25°39'E), collected by A. Smith, 15 January 1973. 

Note standard form of writing South African Museum registration numbers and date. 

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Capital initial letters 

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Biological Abstracts. 



E. B. EASTWOOD 

NOTES ON THE SCORPION FAUNA OF THE CAPE 

PART 3 
SOME OBSERVATIONS ON THE DISTRIBUTION 

AND BIOLOGY OF SCORPIONS ON 

TABLE MOUNTAIN 



VOLUME 74 PART 11 JANUARY 1978 



ISSN 0303-2515 



MUS. COMP. ZOOL-. 
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Bullough, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan. 

Fischer, P.-H. 1948. Donnees sur la resistance et de le vitalite des mollusques. /. Conch., Paris 88: 100-140. 

Fischer, P.-H., Duval, M. & Raffy, A. 1933. Etudes sur les echanges respiratoires des littorines. Archs 

Zool. exp. gen. 74: 627-634. 
Kohn, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon. 

Ann. Mag. not. Hist. (13) 2: 309-320. 
Kohn, A. J. 19606. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean. 

Bull. Bingham oceanogr. Coll. 17 (4): 1-51. 
Thtele, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. In: Schultze, L. Zoologische 

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ANNALS OF THE SOUTH AFRICAN MUSEUM 
ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM 



Volume 74 Band 
January 1978 Januarie 
Part 11 Deel 




NOTES ON THE SCORPION FAUNA OF THE CAPE 

PART 4 

THE BURROWING ACTIVITIES OF SOME 

SCORPIONIDS AND BUTHIDS 

(ARACHNIDA, SCORPIONIDA) 

By 
E. B. EASTWOOD 



Cape Town Kaapstad 



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NOTES ON THE SCORPION FAUNA OF THE CAPE 

PART 4 

THE BURROWING ACTIVITIES OF SOME 
SCORPIONIDS AND BUTHIDS (ARACHNIDA, SCORPIONIDA) 

By 

E. B. Eastwood 

South African Museum, Cape Town 

(With 1 figure and 2 tables) 

[MS. accepted 23 August 1977] 

ABSTRACT 

Burrowing activities of the scorpionid Opisthophthalmus macer Thorell were studied in 
the field and laboratory, and comparisons made with O. capensis (Herbst). O. karrooensis 
Purcell, O. pallidipes Koch and O. peringueyi Purcell were found to burrow only in their 
immature stages. The morphological adaptions of the adults are discussed in relation to their 
scrape-dwelling habit. The specialized digging behaviour of the buthid Parabuthus planicauda 
(Pocock) is described. Pregnant females were often found to construct proper burrows prior 
to parturition. 

CONTENTS 

PAGE 

Introduction 249 

Discussion 250 

Conclusions 253 

Acknowledgements 254 

References 254 

INTRODUCTION 

Opisthophthalmus macer Thorell is a true burrowing scorpionid, distributed 
from False Bay to Port Elizabeth along the south coast. In this paper the burrow 
morphology and burrowing behaviour of this scorpion are reported and com- 
pared to the corresponding activities of O. capensis (Herbst). 

O. karrooensis Purcell, O. pallidipes Koch and O. peringueyi Purcell all 
occupy the drier semi-desert regions of the Cape, namely the Karoo and 
Namaqualand southwards to Piketberg. The young of these species are true 
burrowers, whereas the adults have in common a scrape-dwelling habit with 
special morphological adaptions for this particular way of life. 

Digging behaviour has been reported in the Buthidae and shows certain 
similarities to Parabuthus planicauda (Pocock) and P. capensis (Ehrenberg). 
This was illustrated by Pavlovsky (1924) for the transcaspian species Liobuthus 
kessleri Birula, where the first three pairs of walking legs were used for excava- 
tion. Lankester (1882) reported on the digging behaviour of the Egyptian 
species Buthus australis citrina (Hemprich & Ehrenberg). In this case the 

249 
Ann. S. Afr. Mus. 74 (11), 1978: 249-255, 1 fig., 2 tables. 



250 ANNALS OF THE SOUTH AFRICAN MUSEUM 

pedipalps aided the first three pairs of legs in digging operations. 

P. planicauda is a widely distributed buthid of the southern Cape, extending 
from Worcester and Tulbagh in the west to Albany and Cathcart in the east 
(Lawrence 1955). Under certain conditions this species may construct true 
burrows. The specialized digging behaviour of this species is discussed in 
relation to the activities of P. capensis (Ehrenberg). 

DISCUSSION 

Opisthophthalmus macer Thorell 

Two demes of this species were studied. The first was found on the southern 
slopes of the Kleinriviersberge in a habitat consisting of macchia, namely 
Proteaceae, Restionaceae, Ericaceae, Geraniaceae and other families. Available 
ground cover was sandstone and quartzite debris and the substratum was deep, 
sandy loam rich in organic matter. 

O. macer was found to burrow only beneath suitable ground cover. 
Typically a run was constructed with a single entrance, leading to the burrow 
opening. Often an antechamber was present, the probable function of this 
being a feeding area. The burrows of females were invariably longer than those 
of males and often males were found to construct only a run. Table 1 gives the 
measurements of twenty burrows and runs. 

The second deme was located next to the sea in the southern part of 
Walker Bay. This habitat consisted of rocky limestone slopes with very little 
soil and low coastal scrub. In this habitat the burrows were generally shorter 
and none were found with an antechamber in the run. Specimens were also more 
darkly coloured. Table 1 gives the measurements of ten burrows and runs of 
the second deme. 

Burrowing behaviour of O. macer was observed as follows: compacted 
soil was loosened by the action of the chelicerae and scraped into a heap by 
the first two pairs of walking legs. The heap was then moved backward by legs 
I and II, which were tucked beneath the body. Legs III and IV provided the 
traction while the tail was extended, the pedipalps being used as supports. 

The tarsal claws of O. macer are more sharply curved than those of O. capensis. 
According to Newlands (1972) this is related to a pelophilous habit. This view 
could not be substantiated in the present study, since the Kleinriviersberge deme 
was located in sandy loam and the deme from the coastal habitat was found 
to burrow in compacted sand, whereas O. capensis from the Cape Peninsula 
burrowed in sandy loam only. Tail scraping was not observed although the 
heavily sclerotized fifth caudal segment of O. macer is adapted for this function. 

The scrape-dwellers, Opisthophthalmus karrooensis Purcell, O. pallidipes Koch 
and O. peringueyi Purcell 

Observations of a deme of O. karrooensis showed that although the adults 
were found to occupy scrapes or runs, or simply depressions beneath stones, 



NOTES ON THE SCORPION FAUNA OF THE CAPE 



251 



Table 1 

Measurements in centimetres of twenty burrows and runs of Opisthophthalmus macer in 
a habitat of Cape macchia (Deme 1), and ten burrows in a limestone and coastal scrub 

habitat (Deme 2). 



Run length 



Run width 



Burrow entrance height 



Burrow entrance width 



9 
9 



Burrow length <J 

$ 



Vertical depth of burrow 



Deme 1 
Min. Max. Mean 
8,0 19,0 16,5 



Deme 2 
Min. Max. Mean 
6,0 10,0 7,0 



? 



3,0 


14,0 


10,2 


2,5 


7,0 


5,0 


2,5 


3,2 


3,0 


2,2 


3,0 


2,6 


2,5 


3,0 


2,6 


2,2 


2,8 


2,6 


2,5 


3,2 


3,0 








3,0 


3,5 


3,2 








2,0 


3,0 


2,8 


2,0 


3,0 


2,6 


1,8 


2,8 


2,6 


1,4 


2,6 


2,1 


0,0 


12,0 


6,0 


0,0 


6,0 


4,5 


12,0 


18,0 


14,0 


4,0 


8,0 


6,0 


3,0 


8,0 


5,0 


3,6 


6,0 


4,0 


4,0 


11,0 


7,0 


3,0 


10,0 


4,5 



the immature stages constructed burrows characteristic of other groups in 
this genus. These burrows opened beneath ground cover and the length and 
breadth varied with the stage of development. The burrows were often simple, 
straight tunnels varying from 2 to 9 cm in second and third instars to 6 to 17 
cm in subadults. 

The scrape-dwelling habit of adults and burrowing by the young was also 
evident in populations of O. pallidipes and O. peringueyi from the Clanwilliam 
district. Table 2 shows the relationship between the mode of shelter and certain 
morphological characters for ten species of Opisthophthalmus. 

The pedipalps of these three species are powerful and heavily sclerotized. 
The chelae of the males in particular are very long and afford frontal and 
lateral shielding of the opisthosoma (Fig. 1). The mesosoma is broad and 
distinctly dorsoventrally compressed, and the metasoma slightly more laterally 
compressed than those species which burrow in all stages. The tail is adapted 
for tail-scraping operations by being heavily sclerotized. 

Parabuthus planicauda (Pocock) 

While collecting this species in a variety of habitats over a very large area 
it was noticed that pregnant females were usually found in deep, narrow 



252 



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NOTES ON THE SCORPION FAUNA OF THE CAPE 



253 





2 cm 



^ 



Fig. 1 . Pedipalp and prosoma of male specimens of A. Opisthophthalmus karrooensis and 

B. O. macer. 



depressions or properly constructed burrows, opening beneath stones. The 
entrances were oval or round and the burrows varied from 4 to 1 1 cm in length. 
Males, non-pregnant females, and young were found in scrapes of varying 
shapes and lengths. Often specimens occupied natural depressions under stones, 
where there was no evidence of excavation. 

Laboratory specimens of pregnant females were observed scraping sand as 
follows: the tail was curved either sideways or directly over the mesosoma so 
that the telson lay above the carapace, and the first two pairs of walking legs 
were used in alternative scraping motions to loosen the sand. The body was then 
raised by stilting on the hind legs while the first two pairs of legs and sometimes 
the third pair scraped the sand very vigorously so that it sprayed out behind 
the scorpion ; the pedipalps and fourth pair of legs were used as supports during 
this operation. Transport of soil from a burrow was not observed. 

The defensive armature of Parabuthus consists of the well-developed meta- 
soma and highly toxic venom. It was observed that the terminal chambers of 
the burrows were wide enough to enable the sting to be used effectively, and 
the deep narrow depressions were constructed so as to allow the sting to be 
employed without hindrance. 



CONCLUSIONS 

When the burrowing behaviour of O. macer was compared to O. capensis 
only one difference was noted, namely the scraping of soil into a heap by the 
first two pairs of legs prior to removal from the burrow in the former species. 
Eastwood (1978) noted that O. capensis modified its behaviour under different 



254 ANNALS OF THE SOUTH AFRICAN MUSEUM 

substratum conditions. For example, when the soil was dry the scorpion 
transported it backwards out of the burrow in a single rapid movement, whereas 
if it was damp or there were stones or other obstacles in the soil, it was removed 
in a series of backward jerks. Thus it appears that the pattern of behaviour is 
flexible to accommodate various ecological conditions. It may be that the 
scraping of soil into a heap prior to transportation is used by O. capensis, but 
was not observed in this species because the conditions of the soil did not require 
it. However, this observation illustrates that more detailed information is 
required to determine significant specific behaviour patterns. The measurements 
of burrows and runs for the demes occupying two different habitats indicate 
that burrow and run length are a function of the nature of the habitat, i.e. the 
Kleinriviersberge specimens had deeper burrows since the substratum consisted 
of deep, sandy loam, whereas in the coastal habitat of the second deme, 
specimens constructed shallower burrows in pockets of compacted sand on a 
limestone outcrop. Generally O. macer and O. capensis show similar trends 
in burrow construction and behaviour, and more in-depth studies would be 
necessary to determine significant differences. 

The scrape-dwelling species of Opisthophthalmus are adapted to their 
habitat by the development of a flattening of the body and the increased effective- 
ness of defensive and aggressive armature. That the young of these forms are 
burro wers indicates that they may have evolved from the true-burro wers. The 
flattening of the opisthosoma and metasoma and elongation of the chelae are 
also characteristic of the ischnurinine genus Hadogenes, a lithophilous form, 
and is a good example of parallel evolution. 

The burrowing habit of P. planicauda appears to occur only in females 
prior to parturition. The sand-scraping behaviour of this species is essentially 
the same as that reported for P. capensis by Eastwood (1977). Removal of 
soil in large quantities by P. capensis was not observed in P. planicauda. However, 
this process is obviously necessary for burrow excavation, since the confined 
space at the entrance would not allow for the posture required for sand-scraping 
operations. No doubt many other Parabuthus species will be found to burrow 
and a great deal needs to be studied with respect to functional morphology, 
burrowing behaviour and the relationship to the type of habitat. 

ACKNOWLEDGEMENTS 

I thank Dr V. Whitehead for his advice and help, and the Director of 
the South African Museum for providing research and transport facilities. 

REFERENCES 

Eastwood, E. B. 1977. Notes on the scorpion fauna of the Cape. Part 2. The Parabuthus 
capensis (Ehrenberg) species-group; remarks on taxonomy and bionomics (Arachnida, 
Scorpionida, Buthidae). Ann. S. Afr. Mus. 73: 199-214. 



NOTES ON THE SCORPION FAUNA OF THE CAPE 255 

Eastwood, E. B. 1978. Notes on the scorpion fauna of the Cape. Part 3. Some observations 

on the distribution and biology of scorpions on Table Mountain. Ann. S. Afr. Mus. 

74: 229-248. 
Lankester, E. R. 1882. Observations on scorpions. Proc. R. Soc. 33: 95-104. 
Lawrence, R. F. 1955. Solifugae, Scorpions and Pedipalpi, with checklists and keys to South 

African families, genera and species. S. Afr. anim. Life 1 : 1 52-262. 
Newlands, G. 1972. Ecological adaptions of Kruger Park scorpionids (Arachnida: 

Scorpionides). Koedoe 15: 37-48. 
Pavlovsky, E. 1924. Zur ausseren Morphologie der Scorpione. Ezheg. zool. Mus. 25: 125-141. 



6. SYSTEMATIC papers must conform to the International code of zoological nomenclature 
(particularly Articles 22 and 51). 

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Family Nuculanidae 

Nuculana (Lembulus) bicuspidata (Gould, 1 845) 

Figs 14-15A 
Nucula (Leda) bicuspidata Gould, 1845: 37. 
Leda plicifera A. Adams, 1856: 50. 

Laeda bicuspidata Hanley, 1859: 118, pi. 228 (fig. 73). Sowerby, 1871 : pi. 2 (fig. 8a-b). 
Nucula largillierti Philippi, 1861 : 87. 
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Holotype 

SAM- A 13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King's Beach 
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Biological Abstracts. 



E. B. EASTWOOD 

NOTES ON THE SCORPION FAUNA OF THE CAPE 

PART 4 

THE BURROWING ACTIVITIES OF SOME 

SCORPIONIDS AND BUTHIDS 

(ARACHNIDA SCORPIONIDA) 



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Bullough, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan. 

Fischer, P.-H. 1948. Donnees sur la resistance et de le vitalite des mollusques. /. Conch., Paris 88: 100-140. 

Fischer, P.-H., Duval, M. & Raffy, A. 1933. Etudes sur les echanges respiratoires des littorines. Archs 

Zool. exp. gen. 74: 627-634. 
Kohn, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon. 

Ann. Mag. nat. Hist. (13) 2: 309-320. 
Kohn, A. J. 19606. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean. 

Bull. Bingham oceanogr. Coll. 17 (4): 1-51. 
Thtele, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. In: Schultze, L. Zoologische 

und anthropologische Ergebnisse einer Forschungsreise im west lichen und zentralen Sud-Afrika 4 : 269-270. 

Jena: Fischer. Denkschr. med.-naturw. Ges. Jena 16: 269-270. 

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ANNALS OF THE SOUTH AFRICAN MUSEUM 
ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM 



Volume 74 Band 
February 1978 Februarie 
Part 12 Deel 




CRETACEOUS FAUNAS FROM ZULULAND 

AND NATAL, SOUTH AFRICA 
THE AMMONITE FAMILY LYTOCERATIDAE 

NEUMAYR, 1875 

By 

WILLIAM JAMES KENNEDY 

& 
HERBERT CHRISTIAN KLINGER 



Cape Town Kaapstad 



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CRETACEOUS FAUNAS FROM ZULULAND AND NATAL, 

SOUTH AFRICA 
THE AMMONITE FAMILY LYTOCERATIDAE NEUMAYR, 1875 

By 

William James Kennedy 
Geological Collections, University Museum, Oxford 

& 

Herbert Christian Klinger 

South African Museum, Cape Town 

(With 53 figures) 
[MS. accepted 1 September 1977] 

ABSTRACT 

Ammonites referred to the family Lytoceratidae Neumayr, 1875, occur in rocks of 
Barremian, Aptian and Albian age in northern Zululand, South Africa, and represent the 
most southerly Cretaceous records of all the species present. The following are described: 
Lytoceras vogdti Karakasch, which is common in the Aptian, Lytoceras aff. sauclum 
(Drushchitza), represented by a single specimen from the Aptian, and Lytoceras hennigi 
Zwierzycki, which is frequent in the Upper Barremian and was previously known only from 
Tanzania. Eu lytoceras is represented in the Upper Barremian by the widely occurring 
E. phestum (Matheron); A. (Ammonoceratites) mahadeva (Stoliczka) occurs in the Middle and 
Upper Albian, and A. {Ammonoceratites) crenulatum (Crick), A. (A.) ezoense (Yabe) and 
A. (A.) crenocostatum (Whiteaves) are shown to be synonyms of Stoliczka's species. A. 
(Argonauticeras) depereti (Kilian), of which A. (A.) argonautarum (Anderson) and A. (A.) 
belliseptatum (Anthula) are considered synonyms, occurs in the Upper Aptian; a form referred 
to as A. (A.) aff. depereti is present in the Albian, accompanying A. (A.) besairiei Collignon. 
Protetragonites is represented in the Upper Albian by the widely occurring P. aeolus aeolus 
(d'Orbigny). 

The genus Pictetia is regarded as only a doubtful lytoceratid, perhaps better classed with 
the Ancyloceratina ; it is represented by a specimen referred to as Pictetia aff. depressa (Pictet 
& Campiche). A number of important specimens, including the types of A. {Ammonoceratites) 
ezoense, A. {Am.) crenulatum and A. {Argonauticeras) belliseptatum, are figured photographi- 
cally for the first time. 

CONTENTS 

PAGE 

Introduction 257 

Location of specimens 258 

Field localities 258 

Dimensions of specimens . 258 

Suture terminology 259 

Systematic palaeontology 259 

Acknowledgements 330 

References 330 

INTRODUCTION 

The Lytoceratidae are represented by nine species in the South African 
Cretaceous, all referrable to the subfamily Lytoceratinae Neumayr, 1875, of 

257 
Ann. S. Afr. Mus. 74 (12), 1978: 257-333, 53 figs. 



258 ANNALS OF THE SOUTH AFRICAN MUSEUM 

which the Hemilytoceratinae Spath, 1927, and Protetragonitidae Spath, 1927, 
are regarded as inseparable parts. 

The Lytoceratinae are the rootstock of the Lytoceratina, ranging from 
Triassic to mid-Cretaceous. They are a morphologically conservative group, 
with long and widely ranging genera and species. During the late Jurassic and 
Cretaceous the group were commonest in the Mesogean Realm, although known 
from as far north as Greenland and as far south as the sub-Antarctic Islands, 
and typically rare in the Boreal Realm of western Europe and the United 
States Western Interior. 

The following species are described below : 

Lytoceras vogdti Karakasch, 1907 

Lytoceras aff. sauclum (Drushchitza, 1956) 

Lytoceras hennigi Zwierzycki, 1914 

Eulytoceras phestum (Matheron, 1878) 

Ammonoceratites {Ammonoceratites) mahadeva (Stoliczka, 1865) 

Ammonoceratites {Argonauticeras) depereti (Kilian, 1892) 

Ammonoceratites {Argonauticeras) aff. depereti (Kilian, 1892) 

Ammonoceratites {Argonauticeras) besairiei Collignon, 1949 

Protetragonites aeolus aeolus d'Orbigny, 1850 

Pictetia aff. depressa (Pictet & Campiche, 1861) 

LOCATION OF SPECIMENS 

The following abbreviations are used to indicate the repositories of the 
materials studied: 

BMNH British Museum (Natural History), London 

IGS Geological Museum, London 

MNHP Museum d'Histoire Naturelle, Paris 

SAM South African Museum, Cape Town 

SAS South African Geological Survey, Pretoria 

GSC Geological Survey of Canada, Ottawa 

UPG University of Pretoria, Geology department 

FIELD LOCALITIES 

Details of field localities referred to in the paper are given in Kennedy & 
Klinger (1975). 

DIMENSIONS OF SPECIMENS 

Dimensions of specimens are given in millimetres; abbreviations are as 
follows : 

D = diameter, Wb = whorl breadth, Wh = whorl height, U = umbilical 
diameter. 

Figures in parentheses are dimensions as a percentage of total diameter. 



CRETACEOUS FAUNAS FROM SOUTH AFRICA 259 

SUTURE TERMINOLOGY 

The suture terminology of Wedekind (1916; see Kullman & Wiedmann 
1970 for a recent review) is followed in the present work: 

I s = Internal lobe with septal lobe, U = Umbilical lobe, L = Lateral 
lobe, E = External lobe. 

SYSTEMATIC PALAEONTOLOGY 

Phylum MOLLUSCA 

Class Cephalopoda Cuvier, 1797 

Subclass ammonoidea Zittel, 1884 

Order LYTOCERATIDA Hyatt, 1899 

Superfamily lytocerataceae Neumayr, 1875 

Family LYTOCERATIDAE Neumayr, 1875 

Subfamily Lytoceratinae Neumayr, 1875 

Genus Lytoceras Suess, 1865 
Types species 

Ammonites fimbriatus J. Sowerby, 1817 (I. C.Z.N. Opinion 130). 

Diagnosis 

Evolute, serpenticone, whorls slowly expanding, with a rounded to quadrate 
whorl section. The surface of the shell is ornamented by fine crenulate ribs 
or growth lines, and there are periodic flares, each associated with a constriction 
on the internal mould. The shell surface may be finely striate. Suture formula 
ELU 2 (U 1V : U X d) I s ; highly subdivided, with a short external lobe (E) and a 
triangular, highly subdivided lateral (L) lobe. There is a large septal lobe. 

Discussion 

Lytoceras is a conservative genus, ranging through the Jurassic and Lower 
Cretaceous with very little change. There have been repeated attempts to sub- 
divide the genus, but the authors follow Arkell & Wright (1957), Wiedmann 
& Dieni (1968) and others in regarding Ophiceras Suess, 1865, Fimbhlytoceras 
Buckman, 1918, Thysanoceras Hyatt, 1867, Thysanolytoceras Buckman, 1905, 
Kallilytoceras Buckman, 1921, Crenilytoceras Buckman, 1926, Orcholytoceras 
Buckman, 1926, Biasaloceras Drushchitza, 1953, Valentolytoceras Beznosov, 
1958, and Dinolytoceras Beznosov, 1958, as synonyms. 

Hemilytoceras Spath, 1927 (type species Ammonites immanis Oppel, 1865), 
has rounded and smooth inner whorls, and on the outer whorl develops high, 
concave, closely spaced lamellar flares. Metalytoceras Spath, 1927 (type species 
Lytoceras triboleti Hohenegger in Uhlig 1883), is a compressed, finely ribbed 
genus, with very weak constrictions, and is readily separable from Lytoceras 
by virtue of the consistent splitting of the ribs into fine riblets over the venter. 
Pterolytoceras Spath, 1927 (type species Ammonites exoticus Oppel, 1865), is 
very loosely coiled, with slowly expanding subcircular whorls; the ornament is 



260 ANNALS OF THE SOUTH AFRICAN MUSEUM 

much finer than in Lytoceras, and rather irregular. 

Eulytoceras Spath, 1927 (type species Ammonites inaequalico status 
d'Orbigny, 1840), diners from Lytoceras in having regular, distant ribs and 
periodic flares, with fine striae or riblets between. In the type species, this 
ornament survives in adults, as it does in E. phestum (Matheron). There are, 
however, species such as ' Biasaloceras' sauclum Drushchitza which have 
Eulytoceras-like inner whorls, but develop a Lytoceras-Wkt adult ornament of 
fine, crinkled ribs and flares with associated constrictions. These forms are both 
classed with Lytoceras sensu stricto here, but they point to the homogeneity of 
the Lytoceratinae and the problem of placing many 'intermediate' species 
within the group. 

The remaining Cretaceous Lytoceratinae are readily distinguished from 
Lytoceras; in Pictetia Uhlig, 1883 (type species Crioceras astierianum d'Orbigny, 
1842), the whorls are not in contact, whilst there are no flares or constrictions. 
These are also generally absent in the rapidly expanding, finely ribbed Ammono- 
ceratites (Ammonoceratites) and A. (Argonauticeras), although, as is noted 
later, the dividing line between many lytoceratid genera discussed here is 
sometimes exceedingly thin. 

Occurrence 

Lytoceras ranges from the Lower Jurassic (Pliensbachian) to the 
Cenomanian, with a world-wide distribution extending as far north as Japan, 
Greenland and Alaska, and as far south as Zululand and the sub-Antarctic 
Islands. In the Cretaceous, the group is unknown in the Boreal region of 
northern Europe and the Western Interior of North America. 

Lytoceras vogdti Karakasch, 1907 

Figs 1-8, 9 A, 12A, F-G 

Lytoceras vogdti Karakasch, 1907: 51, pi. 5 (fig. 1), pi. 24 (fig. 31), pi 26 (fig. 7). Forster, 1975: 

143, pi. 1 (fig. 3). 
Pictetia vogdti Drushchitza, 1956: 82, pi. 5 (fig. 19), text-fig. 36a-d. Drushchitza & 

Kudryavtseva, 1960: 257, pi. 6 (fig. la-b), text-fig. 65. 
'Pictetia' vogdti Schindewolf, 1961 : 679. 

Type 

The larger of the two specimens figured by Karakasch (1907, pi. 5 (fig. 1)) 
is herein designated lectotype of this species; it is from the Barremian of the 
Crimea. 

Material 

Large lytoceratids referred to this species are common in the Upper Aptian 
of Zululand, especially along Mlambongwenya Spruit and in the Mkuze Game 
Reserve; SAS L/Li, Z1113-4, Z805, 805a and A1158, are all from the Makatini 
Formation, Aptian IV, at locality 151 in the Mkuze Game Reserve. SAS Z7 
(Haughton Collection) is from the Makatini Formation at Haughton's (1936) 



CRETACEOUS FAUNAS FROM SOUTH AFRICA 



261 



t 





Fig. 1. Lytoceras vogdti Karakasch, 1907. SAS Z1731. x 0,38. 



locality Z7. UPG B391-393 and BMNH C78716 are from the Makatini 
Formation at locality 171; BMNH C78712, C78715, and C78717 are from the 
same formation, Aptian IV, at locality 172, Mlambongwenya Spruit. SAS 
LJE71, BMNH C78702, and possibly C78710 are from the Makatini Formation, 
Aptian III-IV, at locality 168, Mfongozi Spruit; and BMNH C78703 is from 
the same horizon at locality 34 on a tributary of the Mzinene River near 
Hluhluwe, Zululand. 



262 



ANNALS OF THE SOUTH AFRICAN MUSEUM 




B 



Fig. 2. Lytoceras vogdti Karakasch, 1907. SAS Z1731. x 0,38. 



CRETACEOUS FAUNAS FROM SOUTH AFRICA 



263 




Fig. 3. Lytoceras vogdti Karakasch, 1907. SAS LJE 71. X 0,38. 



264 



ANNALS OF THE SOUTH AFRICAN MUSEUM 




B 



Fig. 4. Lytoceras vogdti Karakasch, 1907. SAS LJE 71. X 0,38. 



CRETACEOUS FAUNAS FROM SOUTH AFRICA 



265 



^X 



i 






, 



' 











Fig. 5. Lytoceras vogdti Karakasch, 1907. A. SAS Z1731, showing septal lobe, x 0,45. B. SAS 
2805, showing relationship between septal lobe and cruciform internal lobe, x 1,3. 



266 



ANNALS OF THE SOUTH AFRICAN MUSEUM 




Fig. 6. External suture of Lytoceras vogdti Karakasch, 1907. x 4,5. 



CRETACEOUS FAUNAS FROM SOUTH AFRICA 



267 




X 




O 



CO 

O 



PQ 



OQ 



Ml 



5 



268 



ANNALS OF THE SOUTH AFRICAN MUSEUM 




B 



Fig. 8. Lytoceras vogdti Karakasch, 1907. Copy of the original figures of the lectotype. x 1. 



Dimensions 

Lectotype (after 
Karakasch) . 


D 

85,0 


Wb 

34,0(40) 


Wh 

32,0(38) 


Wb:Wh 
1,06 


U 

30,0(35) 


SAS L/Li . . 


155,0 


67,5(44) 


67,0(43) 


1,00 


55,0(35) 


SAS Z805 . . 


— 


82,0(-) 


81,0(-) 


1,01 


- (-) 


SASZ805a . . 


— 


76,0(-) 


76,0(-) 


1,00 




UPGB393 . . 


20S,0 


95,0(45) 


94,0(45) 


1,00 


- (-) 


LJE71 . . . 


282,0 


130,0(46) 


135,0(48) 


0,96 


80,0(28) 


SASZ113 . . 


330,0 


149,0(45) 


142,0(43) 


1,05 


- (-) 


UPGB391 . . 


350,0 


165,0(47) 


161,0(46) 


1,02 


115,0(33) 


SASZ1114 . . 


. 405,0 


198,0(49) 


170,0(42) 


1,16 


120,0(30) 


SAS Al 158 . . 


— 


211,0(-) 


200,0(— ) 


1,05 


- (-) 



CRETACEOUS FAUNAS FROM SOUTH AFRICA 269 

Description 

This is a very large lytoceratid ; the largest fragment is still septate at a 
whorl breadth of 211 mm, corresponding to an estimated total diameter of 
close on 450 mm for the phragmocone alone. Almost all specimens have very 
poorly preserved inner whorls, commonly encrusted with oysters, and it has 
proved impossible to dissect out the nuclei of the giant specimens to confirm 
that they indeed correspond to the few associated juveniles, the best preserved 
of which is illustrated as Figures 7A-C, 12A. This individual is partially 
exfoliated and does not preserve the outer shell surface ; it shows, however, very 
evolute coiling with a shallow dorsal impressed zone, a wide umbilicus and 
rapidly expanding depressed whorls (Fig. 7A). The exfoliated ornament (Fig. 
7B-C) consists of prominent, narrow, fairly distant ribs separated by from three 
to four fine growth striae. Both ribs and growth striae arise at the umbilical 
seam; they are concave on the umbilical wall, sweep forwards across the shoulder 
and are markedly prorsiradiate on the flank, crossing the venter with a broad 
shallow convexity (Fig. 7B). The specimen bears a single broad constriction, 
parallel to ribs and striae. 

BMNH C78710 (Fig. 12F-G), a fragment tentatively referred to this species, 
shows perfectly preserved ornament at a whorl height of 42,5 mm. The ribs 
and striae are markedly prorsiradiate and pass across the venter with but a 
shallow convexity. Ribs are distant, and minutely crenulated, whilst between 
them are from three to five growth striae of variable strength and development. 
In addition, there is a delicate spiral ornament connecting the crinkles on 
successive ribs. 

Beyond 150 mm diameter, it is convenient to describe the specimens in 
terms of two morphological extremes, with passage forms between. In variety a 
the dorsum is flattened and the venter quite narrowly rounded. The ribs flex 
slightly backwards over the umbilical wall and are radial to feebly prorsiradiate 
on the flanks, crossing the venter with only slight curvature. There are sparse, 
broad, shallow constrictions bounded by a prominent flared adoral rib. As 
diameter increases, the ribs become widely spaced and in most specimens 
growth striae are not preserved although a few specimens (e.g. Fig. 9A) show 
that both they and the delicate spiral ornament survive. On moulds the ribs 
appear as low, rounded undulations. In variety ft the venter and dorsum are 
equally rounded, and the greatest breadth is at mid-flank— a square cross- 
section in which the corners are rounded. Ribs are distinctly prorsiradiate and 
there are occasional flared ribs, not, apparently, accompanied by constrictions. 

The suture line of this species is deeply and intricately subdivided (Fig. 6); 
L is large and asymmetrically bifid, U 2 larger than U l5 and I cruciform (Fig. 5B). 
The septal lobe is massive (Fig. 5A). 

Discussion 

The lectotype of Lytoceras vogdti is a juvenile, but shows an expansion rate 
and differentiation of ornament into ribs and striae like that of some of the 



270 ANNALS OF THE SOUTH AFRICAN MUSEUM 

present similarly sized fragments, sufficient to suggest them to be conspecific. 
Some of the specimens resemble Lytoceras mikadyense Krenkel (1910: 233, 
pi. 22 (fig. 5)), but this species has consistently wider spaced ribs with striae 
between, indicating it to be closer to Eulytoceras. Of other species described by 
Krenkel, the unique holotype of 'Crioceras' schlosseri Krenkel (1910: 227, 
pi. 22(3) (fig. 15)), although more compressed, has ornament like that of some 
of the present specimens. 

Of species described from approximately contemporaneous strata in 
Madagascar, Lytoceras aff. vicinwn Douville of Collignon (19626: 5, pi. 217 
(fig. 952)) has a compressed whorl section with delicate concave, prorsiradiate 
flank ribs, whilst Lytoceras belliseptatiforme Collignon (19626: 6, pi. 217 
(fig. 953), pi. 218 (fig. 953)) is densely and evenly ornamented by fine ribs, has 
a lower expansion rate and circular whorl section. Lytoceras mahafalense 
Collignon (1963: 14, pi. 246 (fig. 1503)) has a distinctly compressed oval whorl 
section and is of Albian age. The most similar species appears to be Lytoceras 
(Thysanolytoceras) str amber gense mut. albensis Collignon (1963: 9, pi. 244 
(fig. 1047)), a Lower Albian form with rather finer ribbing and a distinctively 
compressed whorl section and lower expansion rate. The species may be a 
descendant of the present form. 

Occurrence 

Lytoceras vogdti was first described from the Barremian of the Crimea; 
it has been subsequently recorded from the Upper Aptian of Mozambique, 
and now Zululand. 

Lytoceras aff. sauclum (Drushchitza, 1956) 

Figs9B, 10A-D, 11B 
Compare : 

Biasaloceras sauclum Drushchitza, 1956: 71, pi. 4 (fig. 13); Drushchitza & 

Kudryavtseva 1960: 256, pi. 5 (fig. 1). 
Lytoceras sauclum Wiedmann and Dieni, 1968: 29. 

Type 

The holotype is Drushchitza's (1956, pi. 4 (fig. 13)) original specimen from 
the Hauterivian of the Crimea. 

Material 

BMNH C78704 from the Makatini Formation, Albian IV, locality 171, 
Mlambongwenya Spruit, Zululand. 

Dimensions 

D Wb Wh Wb:Wh U 

BMNHC78704 . 122,0 52,5(43) 48,0(39) 1,1 49,0(40) 



CRETACEOUS FAUNAS FROM SOUTH AFRICA 



271 




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272 ANNALS OF THE SOUTH AFRICAN MUSEUM 

Description 

The coiling is very evolute, serpenticone, with a narrow, shallow, impressed 
dorsal area. The whorl section is depressed (whorl breadth to height ratio may 
be up to 1,1), expanding quite rapidly, with the greatest breadth some way 
below mid-flank. The umbilical seam is deeply incised, the umbilicus of moderate 
width (about 40% of diameter) and depth. The umbilical wall is high and 
rounded, the umbilical shoulder broadly rounded, the flanks flattened, ventro- 
lateral shoulders broadly rounded, and the venter somewhat flattened. 

The shell is ornamented by growth striae, and widely spaced crinkled ribs. 
Ribs and striae arise at the umbilical seam, curve backwards across the umbilical 
wall and forwards across the umbilical shoulder and are straight and slightly 
prorsiradiate across the flanks. The inner whorls have a distinctive Eulytoceras- 
like appearance, with between seven and ten striae between each pair of ribs, 
the ribs themselves being distinctly— if minutely— crenulate, with a gently 
inclined apical slope and an abrupt apertural face which give the surface of the 
shell a scale-like appearance (Fig. 9B). As size increases (Fig. 10) the ribs become 
more closely spaced, striae lose their prominence, crenulations become more 
distinct, and on some specimens, fine strigations appear on some areas of shell. 
All specimens bear occasional broad, deep constrictions on both shell and 
mould, preceded by a high, flared rib. Both flares and constrictions become 
increasingly prominent as diameter increases. 

The suture line (Fig. 1 IB) includes a deeply incised elongate E with a 
long, narrow lanceolate median saddle, a large, triangular, asymmetrically 
bifid L, and a similar smaller U 2 lobe, separated by a large, splayed bifid L/U 2 
saddle. 

Discussion 

Drushchitza figured the type specimen of Biasaloceras sauclum in side view 
only, but the illustration shows clearly the differentiation of ornament on the 
Eulytoceras-like early whorls into ribs with groups of striae between, an inter- 
mediate stage with rather irregular ornament, and middle to late growth 
stages with straight, variably spaced crenulate ribs and occasional flares. The 
whorl section is slightly depressed, with the greatest breadth some way below 
mid-flank. Although of Hauterivian age, the striking similarity between the 
illustration and the present material suggests that they belong to the same 
group, in spite of age difference. 

Biasaloceras striatum Drushchitza (1956: 78, pi. 4 (figs 14a-b); Drushchitza 
& Kudryavtseva 1960, pi. 4 (figs 5a-b)) has a similar juvenile ornament, but a 
rather different whorl section; based on a juvenile, the authors are inclined to 
regard it as a possible variant of L. sauclum. The markedly different early and 
late ornament of Lytoceras sauclum helps to differentiate it from most Cretaceous 
Lytoceras species. Thus, of the long-ranging Stramberg species, Lytoceras 
liebigi Oppel (Zittel, 1868: 74, pi. 9 (figs 5a-b, 7a-b), pi. 10) has a depressed 
oval whorl with concave crenulate ribs. Lytoceras strambergense Zittel (1868: 



CRETACEOUS FAUNAS FROM SOUTH AFRICA 



273 




x 

© 
r- 

oo 
r- 

U 

X 

z 

pq 






N 

43 



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s 

3 



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E 



274 ANNALS OF THE SOUTH AFRICAN MUSEUM 

74, pi. 11 (figs 1-3)) has flexuous crenulate ribs, and Lytoceras sutile Oppel 
(1865: 551; Zittel 1868: 76, pi. 12 (figs 1-5)) has a circular whorl section and 
flexuous ribbing throughout ontogeny. Lytoceras aff. strambergense of Collignon 
(1949a: 71, pi. 12 (fig. 1), text-fig. 5) from the Hauterivian of Madagascar has 
distinctly concave ribs and flares. Lytoceras mahafalense Collignon (1963: 14, 
pi. 246 (fig. 1053)) from the Albian of Madagascar has a distinctly compressed 
whorl section and flattened flanks, whilst Lytoceras vicinum Douville (1916: 
93, pi. 1 1 (fig. 6)) is also a compressed species with an elliptical to oval whorl 
section. A final species to mention, Lytoceras hennigi Zwierzycki, described 
below, has a depressed oval whorl section, and a very even, regular ornament. 

Occurrence 

Lytoceras sauclum was originally described from the Hauterivian of the 
Crimea and is also recorded from the Valanginian of Sardinia. 

Lytoceras hennigi Zwierzycki, 1914 

Figs 11A, 12B-E, 13A-E 

Lytoceras sp. Krenkel, 1910: 224, pi. 22 (fig. 10). 

Lytoceras hennigi Zwierzycki, 1914: 40, pi. 4 (figs 6-7). Spath, 1939: 7. 

Type 

Lectotype herein designated is Zwierzycki's (1914, pi. 4 (figs 6-7)) original 
figured specimen from the Trigonia schwartzi Bed (Barremian-Aptian) of 
Mikadi, Tanzania. 

Material 

Fragments of five individuals and one fairly complete specimen, BMNH 
C78705-9, C78711, from the Makatini Formation, Barremian 1, locality 171, 
Mlambongwenya Spruit, northern Zululand. 

Dimensions 

D Wb Wh Wb:Wh U 

Lectotype (from 

Zwierzycki) . . 155,0 58,0(37) 57,0(37) 1,02 61,0(39) 

BMNH C78708 . — 
BMNH C78705 . — 

Description 

The coiling is very evolute, with a narrow, shallow impressed area, the 
whorls expanding quite rapidly. The whorl section is a depressed oval, the 
greatest breadth being at mid-flank. Ornament consists of fine ribs, growth 
striae, and occasional flares. These arise at the umbilical seam, sweep sharply 
backwards over the inner umbilical wall where they are concave, then forwards 



58,0(37) 


57,0(37) 


1,02 


37,0(-) 


31,0(-) 


1,19 


35,0( ) 


31,8( ) 


1,10 



CRETACEOUS FAUNAS FROM SOUTH AFRICA 



275 





Fig. 11. A. Partial suture of Lytoceras hennigi Zwierzycki, 1914. BMNH C78705. x 5. 
B. Partial suture of Lytoceras aff. sauclum (Drushchitza, 1956). BMNH C78704. X 4,5. 



276 ANNALS OF THE SOUTH AFRICAN MUSEUM 

to pass straight across the umbilical shoulder and flanks where they are markedly 
prorsiradiate, to cross the venter with a faint forwards projection. The best 
preserved specimens show the ribs to have been even, sharp-topped and 
minutely crinkled, and separated by wider interspaces ornamented by irregular 
fine striae. Delicate spiral ridges connecting crinkles are present over the 
whole of the shell surface when well preserved. There are periodic high, sharp 
flares (Fig. 12D-E), although these are only rarely preserved. When shells 
are partially exfoliated, growth striae, crinkles and strigations disappear, 
leaving only evenly spaced, rounded ribs and occasional constrictions (corres- 
ponding to the site of flares). Moulds are smooth save for constrictions 
(Fig. 12D). 

The sutures are poorly exposed, but include a large E/L and smaller L/U 2 , 
both deeply incised, asymmetric and triangular, separated by a deep splayed 
L (Fig. 11 A). A distinct siphonal band is present in one specimen (Fig. 12D). 

Discussion 

The present specimens are much smaller than Zwierzycki's lectotype, but a 
comparison with topotype specimens in the British Museum (Natural History) 
suggests that they are indeed referrable to Lytoceras hennigi. The whorl section 
of topotype specimens is depressed at small diameters, whilst the ornament 
consists of even, crinkled ribs with smooth, somewhat wider interspaces between, 
and occasional strong periodic flares, marked by broad constrictions on the 
internal mould. 

The regular, fine ribs and periodic flares of L. hennigi readily distinguish 
it from Eulytoceras species discussed below, whilst Lytoceras liebigi, L. stramber- 
gense and L. sutile differ in proportions and ornament, as noted above. Lytoceras 
subsequens Karakash (1907: 49, pi. 5 (fig. 9, 9a), pi. 24 (fig. 32)) lacks prominent 
constrictions and flares; L. afT. sauclum has completely different, Eulytoceras- 
like juvenile ornament as discussed above, whilst Lytoceras puezanus Haug 
(1889: 197, pi. 8 (fig. 1) pi. 10 (fig. 1)) has less marked flares and very strong 
strigations. Lytoceras densefimbriatum Uhlig (1883: 191, pi. 6 (figs la-c, 2)) is 
a very finely ornamented, compressed form with weaker flares and stronger 
strigations, Lytoceras mahafalense Collignon is compressed, whilst Lytoceras 
subfimbriatum d'Orbigny (1840: 121, pi. 35) has a compressed whorl section, 
very fine fimbriate ornament, and distant flares, four to five per whorl. 

Occurrence 

Lytoceras hennigi is known from the Lower Cretaceous of Tanzania and 
the Upper Barremian of Zululand only. 

Genus Eulytoceras Spath, 1927 
Type species 

Ammonites inaequalicostatus d'Orbigny 1840. 



CRETACEOUS FAUNAS FROM SOUTH AFRICA 



277 







Fig. 12, A, F, G. Lytoceras vogdti Karakasch, 1907. A. BMNH C78703. F-G. BMNH C78710. 
X 1. B-E, Lytoceras hennigi Zwierzycki, 1914. B-D. BMNH C78708. E. BMNH C78705. 

x 1. 



278 



ANNALS OF THE SOUTH AFRICAN MUSEUM 





t 





Fig. 13. Lytoceras hennigi Zwierzycki, 1914. A-B. BMNH C78705. C-E. BMNH C78706. 

x 1. 



CRETACEOUS FAUNAS FROM SOUTH AFRICA 279 

Diagnosis 

Compressed to round-whorled serpenticone lytoceratids with slowly 
expanding whorls ornamented by fine, regular, distant rectiradiate to prorsi- 
radiate ribs and periodic flares, all of which extend across the venter without 
interruption. 

Discussion 

The regular, fine, distant ribs and occasional simple flares of typical 
Eulytoceras clearly separate them from Lytoceras itself (with typically crinkled 
ribs and flares and periodic constrictions), Hemilytoceras (with smooth inner 
whorls and high, closely spaced concave flares), Pterolytoceras (with minutely 
crinkled growth lines and irregular fine ribs), Ammonoceratites {Ammonoceratites) 
and A. (Argonauticeras), both of which have fine, dense, subdued ornament. 

As the authors have noted, however, some species such as Lytoceras 
sauclum, described above, have Eulytoceras-like inner whorls, whilst 'Lytoceras' 
mikadyense Krenkel has Eulytoceras inner whorls and develops very closely 
spaced ornament when adult, being transitional to Lytoceras sensu s trie to. 

Occurrence 

Eulytoceras ranges from the Hauterivian to Lower Aptian, and species 
are known from southern Europe, the Crimea, Caucasus, Bulgaria, east Africa 
(Tanzania), South Africa (Zululand), Madagascar, and California. 

Eulytoceras phestum (Matheron, 1878) 
Figs 14A-E, 15 

Ammonites phestus Matheron, 1878, pi. c-20 (fig. 5). 

Lytoceras phestus Uhlig, 1883: 187, pi. 5 (figs 1-4, 20). Trautschold, 1886: 137. Haug, 1889: 
196, pi. 8 (fig. 2). Simionescu, 1898: 59, pi. 2 (figs 4, 10), pi. 20 (fig. 17). Sarasin & 
Schondelmayer, 1901 : 19. Karakasch, 1907: 46, pi. 4 (fig. 10), pi. 20 (figs 5-6). Pervinquiere, 
1907: 64. Kilian & Reboul, 1915: 21, pi. 1 (figs 1-2). Petkovic, 1921: 48. Kulzhinskaya- 
Voronets, 1933: 5, fig. 6. Rouchadze, 1933: 174. Ksiazkiewicz, 1938: 230, pi. 1 (fig. 1). 
Eristavi, 1955: 53; 1957: 59. 

Lytoceras gresslyi Somogyi (non Hantken), 1914, pi. 11 (fig. 12). 

? Lytoceras lorentheyi Somogyi, 1914: 300, pi. 11 (figs 10-11). 

? Protetragonites phestus Nagy, 1967: 62, pi. 2 (fig. 1). 

Eulytoceras phestum Spath, 1927: 64. Drushchitza, 1956: 87, pi. 5 (fig. 18); Drushchitza & 
Kudryavtseva, 1960: 258, pi. 6 (fig. 3). Collignon, 1962a: 95, pi. 214 (fig. 935). Dimitrova, 
1967: 27, pi. 9 (fig. 3). Kotetichvili, 1970: 60, pi. 4 (fig. 3). Vasicek, 1972: 37, pi. 2 (fig. 6). 
Murphy, 1975: 18, pi. 2 (fig. 3). 

Type 

The 'holotype' is Matheron's original specimen, in the collections of the 
Museum d'Histoire Naturelle de Marseille according to Murphy (1975: 18), 
but Matheron in fact gives three figures, and it is not clear if these represent 
individual specimens or composite drawings. Lectotype designation (if necessary) 
must await fuller redescription of the type material. 



280 



ANNALS OF THE SOUTH AFRICAN MUSEUM 



"""V 




<jT 





Fig. 14. Eulytoceras phestum (Matheron, 1878). A-C. BMNH C78698. D-E SAS L7(5). 

A-D, x 1 ; E, x 2. 



CRETACEOUS FAUNAS FROM SOUTH AFRICA 281 



\ 



/ 



/ 




Fig. 15. Partial suture of Eulytoceras phestum (Matheron, 1878) based on SAS L7(5). x 8. 

Material 

Two specimens, SAS L7(5) and BMNH C78698, both from the Makatini 
Formation at locality 170, Mlambongwenya Spruit, Zululand (Barremian I). 

Description 

The material consists of a wholly septate juvenile 27,5 mm in diameter, 
and a body chamber fragment of a larger individual. 

The coiling is very evolute, the whorls only just in contact, with, as a 
consequence, a very narrow impressed area. The coiling is evolute, the umbilicus 
being approximately 30 per cent of the diameter, and shallow with a rounded 
wall. The whorl section is as broad as high, rounded trapezoidal, with a rounded 
ventrolateral shoulder, convergent, somewhat flattened sides, the greatest 
breadth being some way beyond mid-flank, and a broadly rounded venter. 

Inner whorls up to a diameter of 1 5 mm appear to be devoid of all ornament. 
Beyond this ornament consists of fine, narrow ribs separated by wide, smooth 
interspaces. The ribs arise at the umbilical seam, pass forwards across the 
umbilical wall and shoulder and are prorsiradiate and weakly flexed or convex 
on the flanks. Interspaces are ornamented by fine growth striae. 

The external suture is partially exposed, as shown in Figure 15. 

Discussion 

The present specimens match closely with Matheron's original figures in 
terms of general ornament and proportions, his illustrations being based upon 
crushed specimens. Eulytoceras phestum is readily distinguished from Eulytoceras 
inaequalicostatus (d'Orbigny) (1840: 18, pi. 29 (figs 3-4)), for that species 
develops strong flares, nine per whorl in D'Orbigny's figure. Eulytoceras 
raricinctum (Uhlig) (1883:88, pi. 5 (figs 5-7)) is a distantly ribbed form with only 
fifteen ribs per whorl rather than the forty to forty-two typical of E. phestum. 
Eulytoceras rotundatum Drushchitza (1956: 86, pi. 5 (fig. 15)) has a depressed 



282 ANNALS OF THE SOUTH AFRICAN MUSEUM 

whorl section with fewer, more convex ribs. Eulytoceras intemperans (Coquand 
in Matheron) (1878, pi. c-20 (fig. 4-4c)) has only twenty-one ribs per whorl, 
with distinctive striae between, whilst Eulytoceras electra (Coquand in Matheron) 
(1878, pi. c-20 (fig 2a-2b)) has fifteen flexuous ribs per whorl, also with striae 
between. 

The east African 'Lytoceras' mikadyense (Krenkel) (1910: 223, pi. 22 (fig. 5)) 
is a depressed species, has periodic flares, rather irregular ribs, and prominent 
striae between. Eulytoceras belchasifakaense Collignon (1949a: 70, pi. 11 
(fig. 2-2a), text-fig. 4), from the Hauterivian of Madagascar, has straight, 
quite distantly spaced ribs, rather than the slightly flexuous ornament of 
E. phestum, but may subsequently prove to be a synonym. E. ? komihevitraense 
Collignon (1963 : 1 5, pi. 247 (fig. 1055)), from the Middle Albian of Madagascar, 
is said to possess constrictions, and has dense, distinctly concave ribs with a 
ventral sinus. Finally, Eulytoceras lepidum (d'Orbigny) (1840: 149, pi. 48 
(figs 3-4)) bears strong flares, and is more distantly ribbed. 

Occurrence 

Eulytoceras phestum is a typically Barremian species, known from southern 
France, the Tyrol, Czechoslovakia, Hungary, Bulgaria, the Crimea, Georgia 
and the Carpathians, Tunisia, Madagascar, and South Africa. Kilian & Reboul 
(1915: 21) also cite the species from the Lower Aptian of southern France, 
whilst Murphy (1975: 18) records it from the Upper Barremian of California. 

Genus Ammonoceratites Bowdich, 1822 

Subgenus Ammonoceratites Bowdich, 1 822 
Type species 

Ammonoceratites lamarcki Bowdich, 1 822. 

Diagnosis 

Large, evolute, serpenticone lytoceratids with a circular whorl section, 
ornamented by dense, crenulate fine ribs and growth striae, sometimes combined 
with faint spiral striae. Occasional broad, shallow constrictions may appear. 
Suture highly subdivided, with a massive cruciform septal lobe (I s ). 

Discussion 

A general lack of flares and associated constrictions, fine ornament and 
rounded whorls separates Ammonoceratites {Ammonoceratites) from Lytoceras 
Suess, 1865 sensu stricto, whilst lack of regular, distant flares differentiates it 
from Eulytoceras Spath, 1927, Hemilytoceras Spath, 1927, and Pterolytoceras 
Spath, 1927. Ammonoceratites (Argonauticeras) is readily separated by virtue 
of its rapidly expanding whorls and an ornament of very fine, even, generally 
non-crenulate ribs. 

Considerable confusion surrounds the introduction of this genus, as much 



CRETACEOUS FAUNAS FROM SOUTH AFRICA 283 

as that surrounding the type species, as discussed below. Following Casey 
(1960: 2), Ammonoceratites Rafinesque, 1815, is regarded as a nomen nudum 
and Ammonoceras Lamarck, 1822, a nomen dubium (as introduced) over which 
Ammonoceratites Bowdich, 1 822 (both of which have type species based on the 
same specimen), has priority. 

The following species have been referred to the restricted subgenus : 

A. {Ammonoceratites) lamarcki Bowdich (1822: 21, pi. 3 (fig. 14)). 

A. {Ammonoceratites) glossoidea (Lamarck) (1822: 144 in Chenu 1859: 90, 

text-figs 391-392). 
A. {Ammonoceratites) mahadeva (Stoliczka) (1865: 16, pi. 80 (fig. 1— lb)), 

Middle Albian of southern India. 
A. {Ammonoceratites) crenulatum (Crick) (1907: 236), Upper Albian of 

Zululand. 
A. {Ammonoceratites) betiokyense Collignon (1962&: 8, pi. 218 (figs 954-955)), 

Upper Aptian of Madagascar. 
A. {Ammonoceratites) crenocostatum (Whiteaves) (1884: 45, pi. 9 (fig. 2)), 

Albian of British Columbia. 
A. {Ammonoceratites) ezoense (Yabe) (1903: 9, pi. 1 (fig. 1), pi. 5 (fig. 1)), Upper 

Albian of Japan. 

Occurrence 

Ammonoceratites ranges from Upper Aptian to Upper Albian. There are 
records from British Columbia, southern France, southern India, Japan, 
Madagascar, and South Africa. 

Ammonoceratites {Ammonoceratites) mahadeva (Stoliczka, 1865) 

Figs 16-33 

? Ammonoceratites lamarcki Bowdich, 1822: 21, pi. 3 (fig. 14). 

? Ammonoceras glossoidea Lamarck, 1 822 : 644. 

? Ammonoceratites glossoidea Chenu, 1859: 90, text-figs 391-392. 

Ammonites mahadeva Stoliczka, 1865: 165, pi. 80 (fig. 1). 

Ammonites crenocostatus Whiteaves, 1876: 45, pi. 9 (fig. 2). 

Lytoceras batesi Whiteaves (non Trask), 1884: 202, pi. 27 (fig. 1); 1900: 270. 

Lytoceras mahadeva Kossmat, 1895: 17. 

? Lytoceras ezoense Yabe, 1903: 9, pi. 1 (fig. 1), pi. 5 (fig. 1). 

? Lytoceras crenulatum Crick, 1907: 236. 

? Lytoceras sp. aff. mahadeva: Jeannet, 1908: 105-119, pis 3-6. 

Lytoceras {Ammonoceratites) crenulatum: Breistroffer, 1936: 169, text-fig. lOh. 

Ammonoceratites cf. mahadeva Collignon, 19496: 45, text-fig. 6. 

Ammonoceratites mahadeva: Collignon, 1963: 12, pi. 245 (fig. 1051). 

Ammonoceratites crenocostatus McLearn, 1972: 22, pi. 1 (fig. 5), pi. 2 (figs 1-3). 

Type 

The holotype is Stoliczka's (1865, pi. 80 (fig. 1)) original specimen from 
the Lower Utatur Group of the environs of Moraviatoor, southern India, and 
presumably of Albian age. The original illustrations are reproduced here as 
Figures 27-28. 



284 



ANNALS OF THE SOUTH AFRICAN MUSEUM 



Material 

Eight specimens: BMNH CI 8265, the holotype of Lytoceras crenulatum 
Crick, from the Albian of the Mzinene River area; BMNH C78890, from Bed 
2 of the Mzinene Formation at locality 35 on the Mzinene River (Albian III); 
BMNH C78700-1, from the Mzinene Formation at locality 54 on the Mzinene 
River (Albian V), SAS Al 195 and 2004 come from the same horizon at locality 
53. SAS Z418 and Z428 in the Van Hoepen Collection are from Impala, Ndumu, 
Mzinene Formation (Albian II or III). 



Dimensions 

Holotype (after 

Stoliczka . 
A. ezoense (after 

Yabe) .... 
A. crenocostatum 

holotype, GSC 

4987 (after 

McLearn) 
A. crenocostatum, 

GSC 4976 . . 
BMNH C78700 

SAS Al 195, at . 
SAS Z428, at . 

at 
BMNH C78890 
SAS Z418 . . 
SASA2008 . . 



D 



218,0 



200,0 



Wb 
82,0(38) 
75,0(37) 



Wh 



Wb:Wh 



78,0(36) 1,05 



U 



95,0(44) 



80,0(40) 0,94 70,0(35) 



44,0 



14,7(33,5) 15,4(35,0) 0,95 



115,0 
137,0 
194,0 
220,0 

253,0 



47,0(— ) 
44,8(— ) 
29,0(— ) 
45,0(39) 
49,0(36) 
75,0(39) 
93,0(42) 
78,0(— ) 
98,0(39) 



40,0(-) 
41,5(-) 
28,0(— ) 
44,0(38) 
48,0(36) 
74,0(38) 
88,0(40) 
76,0(-) 
96,0(38) 



1,08 
1,08 
1,04 
1,03 
1,02 
1,03 
1,05 
1,03 
1,01 



20,7(47) 

- (-) 

- (-) 

- (-) 
48,0(42) 
58,0(42) 
83,5(43) 
85,0(38) 

- (-) 
101,0(40) 



Description 

Very large (the Zululand specimens are still septate at diameters of over 
300 mm), very evolute, the whorls only just in contact with a narrow, shallow 
dorsal impressed zone. The whorl section is almost circular, typically just 
slightly broader than high, the ratio varying from 1,0 to 1,05. The whorls expand 
quite rapidly and the umbilicus is broad, varying from 38 to 42 per cent of the 
diameter and increasing slightly during ontogeny, with a deeply incised umbilical 
seam. 

Ornament changes markedly throughout growth. At the smallest diameters 
visible (Figs 16 A, 31C) the shell surface appears virtually smooth, bearing 
only fine prorsiradiate growth striae, distant, minutely crenulated ribs, and 
occasional flares which correspond to distinct constrictions on internal moulds. 
Ribs, flares and striae arise at the umbilical seam, are concave on the inner 
part of the umbilical wall, then sweep forwards and are prorsiradiate and feebly 



CRETACEOUS FAUNAS FROM SOUTH AFRICA 



285 



•T W 




A 



ii M 



J 








Fig. 16. Ammonoceratites {Ammonoceratites) mahadeva (Stoliczka, 1865). A. SAS A2004. 

B. BMNH C78890. x 1. 



286 



ANNALS OF THE SOUTH AFRICAN MUSEUM 




Fig. 17. Ammonoceratites {Ammonoceratites) mahadeva (Stoliczka, 1865). SAS A2004. x 0,45. 



convex on inner and mid-flank (Figs 17, 20). Beyond this, the crenulated ribs 
become increasingly frequent, and there are sparse prominent flares and 
associated constrictions. In middle and later growth ribs are strong and closely 
spaced (Fig. 20). During these later growth stages, ribs and striae arise at 
the umbilical seam, sweep strongly backwards across the inner part of the 
umbilical wall and forwards over the umbilical shoulder. They are distinctly 
prorsiradiate and variably, although commonly, weakly concave across the 



CRETACEOUS FAUNAS FROM SOUTH AFRICA 



287 




Fig. 18. Ammonoceratites (Ammonoceratites) mahadeva (Stoliczka, 1865). SAS A2004. x 0,45. 



flanks and sweep forwards across the ventrolateral shoulder to produce a broad 
ventral peak of variable depth. Growth striae are scarcely visible to the naked 
eye, but the ribs are very distinct and strongly crenulate with a vertical apertural 
face and a gently inclined apical slope. On the ventral and ventrolateral region 
of some of the Zululand specimens there are distinct, curved longitudinal striae 
between and connecting the crinkled ribs, arising from individual crinkles 
(Fig. 20). 

Partially exfoliated shells show low, broad, irregular, fold-like constricted 



288 



ANNALS OF THE SOUTH AFRICAN MUSEUM 




Fig. 19. Ammonoceratites {Ammonoceratites) mahadeva (Stoliczka, 1865). SAS 2418. X 1. 



CRETACEOUS FAUNAS FROM SOUTH AFRICA 



289 






m 




^ -. , M 



•■-.., 



Fig. 20. Ammonoceratites {Ammonoceratites) mahadeva (Stoliczka, 1865). BMNH C78890. 

x 0,6. 



areas during middle and later growth (Figs 22-23), and in some places, transverse 
rows of large pits corresponding to the site of crinkled ribs. Weathered shell 
surfaces are also covered in minute pits. Internal moulds are essentially smooth, 
save for broad, shallow, fold-like constrictions (Fig. 19). 

One specimen shows a dorsal callus, through which traces of the under- 
lying ventral ornament are still visible. 



290 



ANNALS OF THE SOUTH AFRICAN MUSEUM 












vr- 






W 



-.. 



B 




Fig. 21. A. Ammonoceratites {Ammonoceratites) mahadeva (Stoliczka, 1865). BMNH C78890. 
X 0,6. B-D. Ammonoceratites lamarcki Bowdich. Copy of Chenu 1859, text-fig. 391. 



CRETACEOUS FAUNAS FROM SOUTH AFRICA 



291 




Fig. 22. The holotype of Ammonoceratites {Ammonoceratites) crenulatum (Crick, 1907). 

BMNH CI 8265. Slightly reduced. 



The suture line is highly subdivided with a long spatulate ventral saddle, 
a large bifid asymmetric E/L saddle and a smaller also asymmetrically bifid 
L/U 2 saddle, separated by a very large lateral lobe (L), deeper than the external 
lobe (E) and bifid, with a large subdivided median element. \J 1 is highly sub- 
divided, dorsal internal lobe (I) cruciform, septal lobe large (Fig. 18). 

Discussion 

The Zululand specimens show some variation in strength and direction of 
ornament and relative proportions when compared with Stoliczka's magnificent 



292 



ANNALS OF THE SOUTH AFRICAN MUSEUM 




Fig. 23. The holotype of Ammonoceratites {Ammonoceratites) crenulatum (Crick, 1907). 

BMNH CI 8265. Slightly reduced. 



specimen. In particular, the ventral peak is more prominent than shown in 
his figure and the ribs are prorsiradiate rather than radial. The material varies 
in these respects, and rather than trying to separate it into Ammvnoceratites 
mahadeva with straight ribs and a shallow ventral peak and Ammonoceratites 
crenulatum, with prorsiradiate ribs and a deeper peak, the authors regard their 
material as one variable species. It is, however, far from clear whether 
Ammonoceratites mahadeva is the earliest name used for this species. 

Lamarck (1822, vol. 7: 644) introduced the name Ammonoceras glossoidea, 



CRETACEOUS FAUNAS FROM SOUTH AFRICA 



293 




A B C 

Fig. 24. Ammonoceratites {Ammonoceratites) mahadeva (Stoliczka, 1865). BMNH C78700. x 1. 



without figuring the material. His description is brief: 'Cette coquille rompue 
en trois morceaux, qui s'appartiennent successivement, et dont l'un offre 
l'extremite superieure de cette meme coquille, est d'une assez grande taille, 
fort epaisse en sa partie inferieure, arcuee presque en demi-cercle, et se termine 
superieurement en forme de langue. Ses loges sont remplies de matiere pierreuse, 
et leurs cloisons ne se distinguent que dans les parois ou leurs concours forment 
des sutures lobees, laciniees, rameuses, tout a fait analogues a celles des ammo- 
nites. Mais la coquille dont il s'agit en est tres-distincte par sa forme generate; 
car malgre son arcuation, elle n'eut point forme de tours contigus, si la nature 
l'eut agrandie davantage. Sa longueur est de cinquante centimetres.' 

Tn the same year, Bowdich (1822: 21, pi. 3 (fig. 14)) introduced the name 
Ammonoceratites lamarcki, based upon the same fragments. His description is 



294 



ANNALS OF THE SOUTH AFRICAN MUSEUM 




o 



x 

© 
o 
r- 
oo 
r- 
U 

X 

z 



PQ 




GO 



00 



■a 



Xfl 



1 






CRETACEOUS FAUNAS FROM SOUTH AFRICA 



295 




Fig. 26. External suture of Ammonoceratites {Ammonoceratites) mahadeva (Stoliczka, 1865). 

BMNH C78890. x 1,6. 



also brief: 'Septa numerous, undulated at the margins, pi. 3, fig. 14. Siphon 
marginal, interior. Rolled on itself in the same plane ? The locality is unknown, 
M. Lamarck purchased it by accident: he kindly allowed me to take it home 
in order that the figure, the first that has been made, might be as accurate as 
possible.' 

Subsequent authors have varied in their views of the nature of Ammonoceras 
glossoidea— Ammonoceratites lamarcki (the latter has priority ; fide Casey 1960), 
regarding it as of both Jurassic and Cretaceous age and as a species of Hamites, 
Toxoceras, and Pictetia. The age is, however, in all probability Cretaceous, 
for as Chenu (1859: 90) notes, the specimen was not purchased by Lamarck, 
for he (Chenu) quotes Valenciennes as telling him that 'Lamarck lui a souvent 
repete que ce fossile, dont il faisait grand cas, avait ete rapporte par le secretaire 
de la colonie de Pondichery, et donne au fils de Buffon, qui lui en avait luimeme 
fait cadeau. M. Valenciennes ajoute que parmi les ammonites rapportees de 
l'lnde par Jacquemont, il se trouve aussi un fragment de l'A. fimbriatus.' 

Chenu's figure is much better than that of Bowdich, and is reproduced 
here as Figure 21B-D. The specimen was said to be 500 mm in diameter, and 
corresponds to the generally accepted interpretation of Ammonoceratites; its 
occurrence in southern India indicates that it may well be a specimen of what 
has subsequently been termed Ammonoceratites mahadeva. The figures are, 



296 



ANNALS OF THE SOUTH AFRICAN MUSEUM 




W^> 





Fig. 27. Ammonoceratites {Ammonoceratites) mahadeva (Stoliczka, 1861). Copy of Stoliczka's 
figures of the holotype, reduced here by approximately 0,75. 



CRETACEOUS FAUNAS FROM SOUTH AFRICA 



297 




Fig. 28. Ammonoceratites {Ammonoceratites) mahadeva (Stoliczka, 1861). Copy of Stoliczka's 
figure of the holotype, reduced here by approximately 0,75. 

however, so poor that, until the specimen is redescribed and refigured, the 
authors would prefer to use Stoliczka's name, having sought without success 
for the Lamarck/Bowdich type in the Paris museums. 

Of other species, Ammonoceratites {Am.) betiokyense, from the Aptian of 
Madagascar, is based on two small fragments only, but appears to have 
distinctly flattened sides and a markedly asymmetric or even subtrifid U 2 lobe 



298 



ANNALS OF THE SOUTH AFRICAN MUSEUM 










i -a 







!£e\\12 



X 



+ V ' >" 



u 




Fig. 29. Ammonoceratites {Ammonoceratites) ezoense (Yabe, 1903). Lectotype. X 0,75. 



according to Collignon (19626: 8). 

Ammonoceratites {Am.) ezoense (Yabe), lectotype, herein designated, 
Yabe's (1903: 9, pi. 1 (fig. 1), pi. 5 (fig. 1)) original specimen is also an Albian 
species (see range charts in Matsumoto 1954), the original material being 
preserved in the collections of the Geology Department of Tokyo University, 
and refigured here as Figures 29-30, 31A-B. Yabe separated it from Ammo- 



CRETACEOUS FAUNAS FROM SOUTH AFRICA 



299 




Fig. 30. Ammonoceratites {Ammonoceratites) ezoense (Yabe, 1903). Lectotype. x 0,75. 

noceratites {Am.) mahadeva on the basis of more rapidly expanding whorls and 
a smaller umbilicus (Yabe's measurements are reproduced above), but as can 
be seen from the present photographs, the preservation is poor and the specimen 
probably within an acceptable range of variation for A. {Am.) mahadeva as 
shown by the Zululand specimens (e.g. compare Figs 29-30, 31A-B with Figs 
17-19); the authors therefore place it as a questionable synonym. 



300 



ANNALS OF THE SOUTH AFRICAN MUSEUM 




SI 



h *> n I \ 






fiK 




;^V?.f ;^:Hlf/i 
I 






Ji 



B 



* 
* 





Fig. 31 A-B. Ammonoceratites (Ammonoceratites) ezoense (Yabe, 1903). Part of lectotype, 
showing suture. C. Ammonoceratites {Ammonoceratites) mahadeva (Stoliczka, 1861); inner 

whorls of SAS A2004. x 1,5. 



CRETACEOUS FAUNAS FROM SOUTH AFRICA 301 

Ammonoceratites {Am) crenocostatus (Whiteaves) is a further Middle? to 
Upper Albian species known only from the Queen Charlotte Islands off the 
coast of British Columbia. The authors have been able to examine casts of the 
holotype and other specimens from the area, illustrated here as Figures 32-33. 

The holotype is only 43 mm in diameter, and it retains only part of the 
shell. The inner whorls are strongly constricted on the mould up to a diameter 
of 20 mm, with traces of delicate striae where shell is preserved; from 20 to 
40 mm there are distant fine ribs with delicate growth striae between, both 
ribs and striae being prorsiradiate and feebly convex on the venter. Over the 
last preserved part of the specimen the ribs crowd markedly. GSC 4976 is a 
larger specimen, with an original diameter of approximately 120-130 mm. The 
outer whorl shows distinctly concave prorsiradiate flank ribs, with a marked 
convex ventral peak, rib density, form and strength matching closely to the 
similarly sized Zululand specimen BMNH C78700 (Figs 24-25). So far as can 
be judged there are no criteria that can be used to separate A. {Am.) crenocostatum 
from the Zululand material, and it is therefore suggested that it is a further 
synonym of Ammonoceratites {Am) mahadeva. 

Occurrence 

As defined above, Ammonoceratites {Am.) mahadeva ranges from Middle 
to Upper Albian, and is known from southern India, the Queen Charlotte 
Islands, Madagascar, Japan, Zululand, and perhaps the Jura (Jeannet 1908; 
105-119, pis 3-6). 

Subgenus Argonauticeras Anderson, 1938 
Type species 

Lytoceras argonautarum Anderson, 1902. 

Diagnosis 

A subgenus of Ammonoceratites with a high expansion rate, subrectangular 
to trapezoidal whorl section, fine, equal, dense, straight to gently flexed ribs 
or striae without obvious crinkles. Weak, broad constrictions may be present 
on early whorls. 

Discussion 

Argonauticeras can be separated from other Cretaceous lytoceratid genera 
by its fine, even, typically non-crenulate ornament and high expansion rate. 
The trapezoidal whorl section, emphasized by Wright (in Arkell & Wright 
1957), is a less distinctive feature, as discussed below. Differences from 
Ammonoceratites {Ammonoceratites) are outlined above. 

Four species have been referred to the subgenus : 

Argonauticeras argonautarum (Anderson) (1902: 85, pi. 7 (figs 154-155); 
1938: 149, pi. 17 (fig. 3), pi. 19 (figs 1-2)). Upper Aptian of California. 



302 



ANNALS OF THE SOUTH AFRICAN MUSEUM 




o 



* * 




00 



X 

l> 

oo 
On 

u 
o 

Cu 



o 



v© 
oo 



5 



en 
M 

E 



CRETACEOUS FAUNAS FROM SOUTH AFRICA 



303 




X 
r- 

OS 

U 

o 

o 
a 
>. 

X 



so 
oo 



1 




s: 
5 

E 



304 ANNALS OF THE SOUTH AFRICAN MUSEUM 

Argonauticeras depereti (Kilian) (1892: 8, pi. 1 (fig. 2). Thomel, 1968: 684-687, 
pi. 35 (figs 1-4)). Upper Aptian of southern France. 

Argonauticeras besairiei Collignon (1949Z?: 46, pi. 8 (fig. 1)). Upper Aptian to 
Middle Albian of Madagascar. 

Argonauticeras belliseptatum (Anthula) (1899: 97, pi. 6 (fig. 1), pi. 7 (fig. 1)). 
Aptian of the Caucasus and Madagascar. 

Of these species, Dimitrova (1967: 28, pi. 10 (fig. 6)) figured a fragment 
referred to as Pictetia belliseptata (Anthula) which appears actually to uncoil. 
The authors therefore take the opportunity to figure photographically the 
holotype of A. (Ar.) belliseptatum (Figs 40-43) which shows it to be an 
Argonauticeras. 

Pseudotetragonites Drushchitza, 1956 (type species P. kudrjavzevi 
Drushchitza), is a subjective synonym of Argonauticeras, based on juveniles. 

Occurrence 

Argonauticeras is known from the Upper Aptian of southern France, the 
Caucasus, California, the Lower and Middle Albian of Madagascar, and the 
Middle Albian of Zululand. 



Ammonoceratites {Argonauticeras) depereti (Kilian, 1892) 

Figs 34-37, 38B, 39-43 

Lytoceras depereti Kilian, 1892: 8, pi. 1 (fig. 2a-b). Simionescu, 1900: 656. 

Lytoceras belliseptatum Anthula, 1899: 97, pi. 6 (fig. 1), pi. 7 (fig. 1). Collignon, 1963: 12, pi. 

245 (fig. 1052). 
Lytoceras argonautarum Anderson, 1902: 85, pi. 7 (figs 154-155). 
Tetragonites depereti Jacob, 1907: 12. Non Fallot, 1920: 243, pi. 2 (fig. 5), text-fig. 13. 
Lytoceras {Argonauticeras) argonautarum: Anderson, 1938: 140, pi. 17 (fig. 3), pi. 19 (figs 1-2). 
Pseudotetragonites kudrjavzevi Drushchitza, 1956: 80, pi. 7 (fig. 25), text-fig. 35. Orlov, 

1958: 58, pi. 20 (fig. 4), Drushchitza & Kudryavtseva, 1960: 257, pi. 5 (fig. 2a-b), 

text-fig. 64. 
Ammonoceratites {Argonauticeras) depereti Collignon, 1956: 100. Thomel, 1968: 684, pi. 35 

(figs 1-4), text-figs. 
? Pictetia belliseptata Dimitrova, 1967: 28, pi. 10 (fig. 6). 
Argonauticeras belliseptatum Forster, 1975: 144, pi. 1 (fig. 6). 

Type 

The lectotype, herein designated, is Kilian's original specimen from the 
Upper Aptian of the environs of Barreme (Basses Alpes), south-eastern France, 
in the collections of Museum d'Histoire Naturelle, Lyon. 

Material 

Two specimens, UPG B4 and UPG B5 from the Makatini Formation, 
Upper Aptian, Manyola Drift on the Usutu River, at 26°49'15" S 30°13'30" E. 



CRETACEOUS FAUNAS FROM SOUTH AFRICA 



305 




Fig. 34. Ammonoceratites (Argonauticeras) depereti (Kilian, 1892). UPG B4. X 0,78. 



306 



ANNALS OF THE SOUTH AFRICAN MUSEUM 




Fig. 35. External suture of Ammonoceratites (Argonauticeras) depereti (Kilian, 1892). x 0,8. 



Dimensions 

D Wb 

Californian specimen 

(after Anderson, 

1938) .... 250,0 125,0(50) 
Holotype of A. (Ar.) 

belliseptatum (after 

Anthula, 1899) . 325,0 153,0(47) 
Lectotype of A. {Ar.) 

depereti (after 

Kilian, 1892) . . 53,0 25,0(47) 

French specimens 

(after Thomel, 

1968) 

151 ... . 38,3 19,3(50) 

152 . . . . 55,0 27,0(49) 

153 . . . . 102,0 43,0(42) 
UPG B4 . . . . 179,0 70,0(39) 
UPG B5 . . . . 40,0(— ) 



Wh Wb:Wh 



16,0(41) 
23,0(41) 
44,5(43) 
67,0(27) 
37,0(— ) 



1,20 
1,17 
0,96 
1,04 
1,08 



U 



113,0(45) 1,11 87,0(35) 



130,0(40) 1,17 123,0(38) 



27,0(39) 1,19 21,0(39) 



14,4(37) 
19,2(34) 
38,0(36) 
73,0(40) 
- (-) 



Description 

The largest specimen in the collection consists of one and a half whorls, 
parts of which are damaged. Coiling is very evolute, and whorls increase rapidly 
in height and width. The whorl section is slightly wider than high with rounded 
umbilical and ventrolateral edges; the dorsal impressed zone is small. 

Ornament consists of narrow, dense unbranched ribs which pass forwards 
over the flanks with a slight curvature and straight across the venter. Broad, 



CRETACEOUS FAUNAS FROM SOUTH AFRICA 



307 






B 

Fig. 36. Ammonoceratites {Argonauticeras) depereti (Kilian, 1892). UPG B5. x 1. 



low constrictions are also present, paralleling striae ; the number of constrictions 
per whorl cannot be determined, but on the smaller specimen, UPG B5, there 
are four equidistant constrictions in one-third of a whorl. The suture is only 
partially exposed; E is narrow and bifid, L is very wide and deeper than both 
E and U 2 . The saddles E/L and I/U 2 are bifid and have slender stems. 

Discussion 

Thomel recently (1968) reviewed this species and pointed to the changes 
in whorl section, relative dimensions and density of ornamentation during 
ontogeny. The Zululand specimens fall within the range of relative proportions 
given by Thomel (1968: 685) and are virtually identical to the figured specimens 
as far as the whorl section, fine ornamentation and constrictions are concerned. 
The largest known French specimen of A. (Ar.) depereti is only 102 mm in 



308 



ANNALS OF THE SOUTH AFRICAN MUSEUM 





B 




Fig. 37. Ammonoceratites {Ammonoceratites) depereti (Kilian, 1892). A-B. Copies of Kilian's 

figures of the holotype. C. Copy of Anderson's (1938: pi. 17 (fig. 3)) figure of a juvenile 

Lytoceras {Argonauticeras) argonautarum (Anderson, 1902). All figures X 1. 



CRETACEOUS FAUNAS FROM SOUTH AFRICA 



309 




& 






S 



oq 



X 






* 



?3 



be 




310 



ANNALS OF THE SOUTH AFRICAN MUSEUM 




GO 




CRETACEOUS FAUNAS FROM SOUTH AFRICA 



311 




Fig. 40. Ammonoceratites (Argonauticeras) belliseptatum (Anthula, 1899). Holotype. x 0,45. 

(Photograph supplied by R. A. Reyment.) 



312 



ANNALS OF THE SOUTH AFRICAN MUSEUM 




Fig. 41. Ammonoceratites (Argonauticeras) belliseptatum (Anthula, 1899). Holotype. X 0,45. 

(Photograph supplied by R. A. Reyment.) 



CRETACEOUS FAUNAS FROM SOUTH AFRICA 



313 



p 




Fig. 42. Ammonoceratites (Argonauticeras) belliseptatum (Anthula, 1899). Holotype. X 0,45. 

(Photograph supplied by R. A. Reyment.) 



314 



ANNALS OF THE SOUTH AFRICAN MUSEUM 






^*H"i' 









. ;r 0'**. \ 









* A .WlSk 



Fig. 43. Ammonoceratites {Argonauticeras) belliseptatum (Anthula, 1899). Holotype. x 0,45. 

(Photograph supplied by R. A. Reyment.) 



CRETACEOUS FAUNAS FROM SOUTH AFRICA 3 1 5 

diameter, and thus difficult to compare with the holotype of Argonauticeras 
argonautarum (Anderson), the original figures of which are reproduced here 
as Figure 39. A smaller specimen figured subsequently by Anderson (1938: 
pi. 17 (fig. 3)) is reproduced here as Figure 37C; it appears identical to Kilian's 
lectotype (reproduced here as Fig. 37A-B), and in consequence the authors 
regard A. {Ar.) argonautarum as a junior subjective synonym of A. (Ar.) depereti. 
A. {Ar.) belliseptatum Anthula is based on a giant disc 325 mm in diameter. 
When compared with the present material, the overall proportions, expansion 
rate, style and direction of ornament are so similar that the authors believe this 
to be a synonym of A. {Ar.) depereti. The remaining species referred to the 
subgenus, A. {Ar.) besairiei Collignon (1949&: 46, pi. 8 (fig. 1)), is also very 
similar, and, indeed, Forster (1975: 144) regarded it as a synonym of A. {Ar.) 
belliseptatum. The juvenile holotype has, however, a distinctly trapezoidal 
whorl section with a broad, flattened venter, flattened flanks and, according to 
Collignon, has straight and rather coarse ribs for the genus. A larger Mada- 
gascan specimen figured subsequently (Collignon 19626: 10, pis 219-220 (fig. 
956)) has a very high expansion rate, the proportions (after Collignon) being: 



D 


Wb 


Wh 


Wb:Wh 


U 


70,0 


160,0(58) 


136,0(48) 


1,18 


82,0(30) 



Given a larger sample it might prove possible to demonstrate that A. {Ar.) 
besairiei is also within the range of variation of A. {Ar.) depereti, but the authors 
maintain the species separate at this time whilst acknowledging the fine division 
between them. 

Occurrence 

As defined above, Ammonoceratites {Argonauticeras) depereti is known 
from the Upper Aptian of the Caucasus, Bulgaria, southern France, California, 
Mozambique and Zululand, and the Albian of Madagascar. 

Ammonoceratites {Argonauticeras) aff. depereti (Kilian, 1 892) 

Figs 38A, 44-46 
Material 

One specimen only, UPG B465, from the Mzinene Formation, Albian 
III-IV, Mlambongwenya Spruit, northern Zululand. 



Dimensions 














D 


Wb 


Wh 


Wb:Wh 


U 


UPGB465 . . 


47,5 


23,5(49) 


21,0(44) 


1,12 


- (-) 




84,0 


40,0(48) 


38,5(46) 


1,04 


- (-) 




149,0 


68,5(46) 


67,5(45) 


1,02 


- (-) 




248,0 


99,5(40) 


98,5(40) 


1,01 


- (-) 




352,0 


127,5(36) 


134,0(38) 


0,97 


135,0(38) 



316 



ANNALS OF THE SOUTH AFRICAN MUSEUM 




Fig. 44. Ammonoceratites {Argonauticeras) aff. depereti (Kilian, 1892). UPG B465. X 0,5. 



CRETACEOUS FAUNAS FROM SOUTH AFRICA 



317 




Fig. 45. Ammonoceratites {Argonauticeras) aff. depereti (Kilian, 1892). UPG B465. x 0,5. 



Description 

The specimen is beautifully preserved, retaining part of the original shell. 
All but the last quarter whorl are septate. 

Coiling is very evolute with the whorls increasing gradually in height and 
width. The dorsal area of impression is very shallow. The whorl section varies 
with increasing diameter as shown in the table of dimensions. It is initially rect- 
angular, but with increasing diameter the venter becomes narrower and the 
whorl height becomes greater than the whorl breadth. Maximum whorl breadth 
is at the dorsal third of the flanks. The surface is ornamented by numerous thin, 



318 



ANNALS OF THE SOUTH AFRICAN MUSEUM 




Fig. 46. Suture of Ammonoceratites {Argonauticeras) aff. depereti (Kilian, 1892). 

B465. x 3,6. 



/ 



\ 



UPG 



fine ribs. The ribs pass radially over the umbilical wall and then forwards 
across the flanks with a slight curvature, and straight over the venter. On 
the venter of one of the inner whorls, where the specimen had been broken, the 
ribs are minutely crinkled and are connected longitudinally by faint striae. 
At a whorl height of 63 mm there are 45 ribs in a distance equal to the whorl 
height; at 85 mm the figure is 42. On the inner whorls the flanks bear low folds, 
so that the ribs appear to bunch. On the outer whorls this effect is less noticeable, 
but two wide and shallow, barely noticeable constrictions occur. The external 
suture (Fig. 46) is highly divided with narrow-stemmed saddles. The lobes are 
asymmetrically bifid. 

Discussion 

As can be seen from the dimensions, the ratio of Wb/Wh decreases with 
increase in diameter, a situation somewhat comparable to that encountered in 
A. (Ar.) depereti, from which it differs in the lack of constrictions on inner 
whorls (Fig. 38 A), as a consequence of which the authors refer to it as A. (Ar.) 
aff. depereti. It is readily separated from A. {Ar.) besairiei which is markedly 
depressed at large diameters. 

Occurrence 

Middle Albian of northern Zululand. 



Ammonoceratites (Argonauticeras) besairiei Collignon, 1949 

Figs 47-48A 

Argonauticeras besairiei Collignon, 1949a: 46, pi. 8 (fig. 1); 1950: 36, pi. 5 (fig. 6); 19626: 
10, pi. 219-220 (fig. 956). 

Holotype 

The original of Collignon (1949#: pi. 8 (fig. 1)), a juvenile specimen from 
the Lower/Middle Albian transition beds of Ambarimaninga, Madagascar. 



CRETACEOUS FAUNAS FROM SOUTH AFRICA 319 

Material 

A single fragment, BMNH C78697, from the Mzinene Formation, Albian 
III, of locality 171, Mlambongwenya Spruit, northern Zululand. 

Dimensions 





D 


Wb 


Wh 


Wb:Wh 


U 


Holotype (after 












Collignon, 1949a) 


43,0 


21,0(49) 


18,0(42) 


U7 


16,0(37) 


Specimen 1 of 












Collignon, 1950 . 


70,0 


35,0(50) 


33,0(47) 


1,06 


26,0(35) 


Specimen 2 of 












Collignon, 1950 . 


130,0 


51,0(50) 


48,0(47) 


1,06 


28,0(27) 


Original of Collig- 












non, 19626: pis 












219-220 (fig. 956) 


200,0 


106,0(53) 


94,0(47) 


1,13 


63,0(32) 


BMNH C78697 


— 


121,0(-) 


107,0(-) 


1,13 


- (-) 



Description 

The specimen is a wholly septate fragment of a large, massive ammonite 
whose original diameter must have been well over 200 mm. The coiling is very 
evolute, with a narrow, shallow impressed area. The umbilicus is of moderate 
size, and deep. The whorls expand rapidly and are depressed, rounded- 
trapezoidal, the greatest breadth being well below mid-flank. The umbilical 
wall is high, rounded, merging with broadly rounded umbilical shoulders and 
lower flanks. The upper flanks are flattened and converge to a broadly rounded 
venter. 

Ornament consists of fine, non-crenulate, narrow, flattened ribs separated 
by wider interspaces. These arise at the umbilical seam, sweep abruptly back 
across the lower part of the umbilical wall and curve forwards over the shoulder 
to pass across the flanks in a rectiradiate or slightly prorsiradiate direction, 
with a faint convexity. They pass straight across the venter. 

The external suture is not visible, but the fractured sections indicate the 
presence of large L and U 2 lobes, together with a large septal lobe. The dorsum 
shows the impression of the venter of the penultimate whorl, which was 
distinctly flattened. 

Discussion 

At the diameter represented by the Zululand specimen, the trapezoidal 
whorl section of Argonauticeras is lost, but when compared with the series of 
specimens of A. {Argonauticeras) besairiei figured by Collignon (1949^-19626), 
the present specimen is clearly an adult of that species. 

Occurrence 

Upper Aptian to Middle Albian of Madagascar, Middle Albian of 
Zululand. 



320 



ANNALS OF THE SOUTH AFRICAN MUSEUM 



44 







Fig. 47. Ammonoceratites {Argonaut ice ras) besairiei Collignon, 1949. BMNH C78697. X 0,7. 



CRETACEOUS FAUNAS FROM SOUTH AFRICA 321 

Genus Protetragonites Hyatt, 1900 
Type species 

Ammonites quadrisulcatus d'Orbigny, 1841. 

Diagnosis 

Very evolute lytoceratids with circular to oval whorl sections, sparse to 
frequent radial or curved constrictions and a shell ornamented by growth 
striae only. 

Discussion 

Repeated attempts have been made to justify a separation of a family 
Protefragonitidae Spath, 1927, from the Lytoceratidae. This has been based on 
the tendency for species to develop one or more auxiliary elements in the 
suture line. Schindewolf (1960: 681) and Wiedmann (1962: 17) have both 
demonstrated that Protetragonites follow the same sutural development as the 
Lytoceratidae, and are derived from Lytoceras sensu stricto. The authors 
further follow Schindewolf and Wiedmann in regarding Hemitetragonites 
Spath, 1927, and Leptotetragonites Spath, 1927, as subjective synonyms. 

Occurrence 

Protetragonites is a typically Mesogean genus, known chiefly from the 
Mediterranean region, but also known from central Europe (Austria), the 
Crimea, Caucasus, north Africa, Madagascar, and now Zululand. Its strati- 
graphic range is from Tithonian to Upper Albian. 

Protetragonites aeolus aeolus (d'Orbigny, 1850) 
Fig. 48B-F 

Ammonites aeolus d'Orbigny, 1850: 125. 

Lytoceras {Gaudryceras) aeolus Jacob, 1908: 14 (pars), pi. 1 (fig. 17), non pi. 1 (figs 14-16). 

Gaudryceras aeolus: Fallot, 1920: 12. Collignon, 19496: 50. 

Hemitetragonites aeolus Breistroffer, 1936: 175. Breistroffer & Villoutreys, 1953: 71 (pars). 

Protetragonites aeolus aeolus Wiedmann, 1962: 24, pi. 10 (fig. 3), text-fig. 6. 

Protetragonites aeolus: Wiedmann & Dieni, 1968: 31. 

Type 

The lectotype, designated by Wiedmann (1962: 25), is MNHP 5770 in the 
D'Orbigny Collection, Museum d'Histoire Naturelle, Paris, and is from the 
Middle ? Albian of Escragnolles, Var, France. 

Material 

Two specimens, SAS PJR/1-2, from the Mzinene Formation, Albian V, 
at locality 65 along the Munywana Creek, east of Hluhluwe, Zululand, are 
definitely referred to the subspecies, whilst a third specimen, BMNH C78699, 
from the same horizon at locality 56 on the Mzinene River, may also belong 
here. 



322 



ANNALS OF THE SOUTH AFRICAN MUSEUM 



■ - i 




B 




( . ,i 




\ 



\ 




** 



S* 



•Hr 



IK ■ i 



*. 




' 



JfVUtt 




Fig. 48. A. Ammonoceratites(Argonauticeras)besairieiCo\\ignon, 1949. BMNH C78697. x 0,7. 
B-F. Protetragonites aeolus aeolus (d'Orbigny). B-C. PJR/1. D-F. BMNH C78699. B-C, X 1; 

D, x 2; E-F, x 1. 



CRETACEOUS FAUNAS FROM SOUTH AFRICA 323 



Dimensions 














D 


Wb 


Wh 


Wb:Wh 


U 


Lectotype, MNHP 












5770 .... 


31,0 


12,5(40) 


10,5(34) 


1,19 


14,5(47) 


PJR/1 .... 


30,7 


12,5(41) 


8,7(28) 


1,43 


14,3(47) 


BMNH C78699 












(approximate) 


30,1 


- (-) 


9,3(31) 


— 


15,0(50) 



Description 

The coiling is very evolute, serpenticone, with a very wide umbilicus 
(47-50 % of diameter). The whorl section is depressed, reniform, with a gently 
rounded venter and swollen flanks (the greatest breadth is below mid-flank). 
The dorsal impressed zone is narrow and shallow. 

All specimens are corroded and no external shell surface survives except 
in a few areas which retain a suggestion of growth striae. There are four straight, 
prorsiradiate constrictions per whorl, bounded aperturally by a thickened rib. 

The sutures are undecipherable. 

Discussion 

Relative proportions, expansion rate, position and form of constrictions 
indicate these specimens to be Protetragonites, most closely resembling the 
well-known species Protetragonites aeolus (d'Orbigny). Wiedmann (1962: 23) 
recognized three subspecies, P. aeolus aeolus, P. aeolus neptuni Wiedmann 
(1962: 29, pi. 1 (fig. 2), text-fig. 7b), and P. aeolus aeoliformis (P. Fallot) 
(Wiedmann 1962: 26, pi. 1 (fig. 1), pi. 2 (fig. 4), text-fig. 4a). P. aeolus aeoliformis 
has a more or less rounded whorl section rather than reniform as in the nominate 
subspecies, whilst P. aeolus neptuni has strongly ornamented inner whorls and 
much more prominent constrictions. 

Of other species, Protetragonites crebrisulcatus (Uhlig) (Wiedmann 1962: 
19, pi. 1 (fig. 3), pi. 3 (figs 2, 4)) has rather similar overall proportions, but 
bears more constrictions which efface at maturity. P. obliquestrangulatus 
(Kilian) and its subspecies (Wiedmann 1926: 21 et seq.) bear many more prorsi- 
radiate constrictions, whilst Protetragonites laevis Wiedmann (1962: 31, pi. 10 
(fig. 4) text-figs 9-10) has a subquadrate depressed whorl section and is smooth. 

Occurrence 

Protetragonites aeolus and its subspecies are best known from the western 
Mediterranean, i.e. the Balearics, Sardinia and south-eastern France, but there 
are also records from Madagascar, and now Zululand. The species is restricted 
to the Albian. 

SYSTEMATIC POSITION UNCERTAIN 

Genus Pictetia Uhlig, 1883 
Type species 

Crioceras astierianus d'Orbigny, 1842. 



324 ANNALS OF THE SOUTH AFRICAN MUSEUM 

Diagnosis 

Loosely coiled with whorls separated throughout; body chamber may 
straighten. Whorl section rounded, compressed to depressed; ornamented by 
weak, irregular, feebly crinkled ribs. 

Discussion 

Pictetia is an enigmatic genus, known chiefly from septate fragments only 
which are either curved or straight. Few good illustrations of the type species 
are available and in consequence the authors reillustrated (Fig. 49) the original 
of D'Orbigny (1842: pi. 115 (figs 3-5)) which is herein designated lectotype of 
P. astieriana (the specimen has been commonly referred to as the holotype, but 
D'Orbigny clearly states that he was aware of specimens in the Astier, Duval 
and Puzos collections, although that which he figured was 'a leve tous mes doutes 
a ete decouverte par M. Astier'. The authors also take the opportunity of 
illustrating two of the straight fragments, doubtful paralectotypes, also from 
Escragnolles, Var (Figs 50-51). 

In his generic diagnosis of Pictetia, Casey (1960: 3) indicated that it had a 
'suture line with simple lobe formula I U L E, but highly complex'. Such a 
formula is not typical of the Lytoceratina, which are characterized by a quin- 
quelobate suture. Unfortunately there is no detailed work on the sutural 
ontogeny of Pictetia', Schindewolf (1961 : 678) examined specimens of the type 
species, and concluded that the small incision in saddle I/U should be regarded 
as an incision, and not a lobe. If this be true, then Pictetia is either to be trans- 
ferred to the Ancyloceratina (some members of which have fimbriate ornament), 
as did Schindewolf and Hyatt (1900: 588) before him, or retained within the 
Lytoceratida as an uncoiled Lytoceras (Casey 1960: 3) in which the quadrilobate 
suture is secondarily derived from quinquelobate ancestors. In Spiroceras 
bifurcati (Quenstedt) Schindewolf (1961: 762, text-fig. 45) demonstrated that 
two of the original five lobes became so reduced that a pseudo-trilobate condition 
prevailed in adults; might not reduction in Pictetia be a similar reflection of 
acquisition of the heteromorph habit ? Unfortunately the Zululand material does 
not help in the resolution of this interesting problem. 

Occurrence 

Pictetia is known from the Lower and Middle Albian of western Europe, 
Transcaspia, India, Madagascar and Zululand; Collignon (1962&: 13) has 
recorded two species from the Upper Aptian of Madagascar. 

Pictetia aff. depressa (Pictet & Campiche, 1861) 
Figs 52A-C, 53 
Compare : 

Crioceras depressus Pictet & Campiche, 1861 : 28, pi. 45 (figs 3-4). 
Pictetia depressa Casey, 1960: 5, pi. 1 (fig. la-c), text-fig. la-d (with synonomy). 
Collignon, 1962b: 13, pi. 221 (fig. 457). 



CRETACEOUS FAUNAS FROM SOUTH AFRICA 



325 





O 



x 

CO 

m 

o\ 

X 

z 

pq 

<S 



o 

oo 



a 

O 
3 

Q 

K 

a 



ON 

E 



326 



ANNALS OF THE SOUTH AFRICAN MUSEUM 




* 




Fig. 50. Pictetia astieriana (d'Orbigny, 1842). BMNH C5440, wholly septate ancyloceratid- 

like shaft, x 1. 



CRETACEOUS FAUNAS FROM SOUTH AFRICA 



327 





B 



Fig. 51. Pictetia astieriana (d'Orbigny, 1842). BMNH C5439, septate straight shaft. X 1, 



328 



ANNALS OF THE SOUTH AFRICAN MUSEUM 






B 






Fig. 52. A-C. Pictetia aff. depressa (Pictet & Campiche, 1861). SAS LJE 173. x 1. 
D-F. Pictetia depressa (Pictet & Campiche, 1861). Copies of Pictet & Campiche's original 

figures of the species. 



CRETACEOUS FAUNAS FROM SOUTH AFRICA 329 




Fig. 53. Pictetia aff. depressa (Pictet & Campiche, 1861). Suture of SAS LJE 173. x 3,6. 

Material 

One specimen only, SAS LJE 173 from the Mzinene Formation, Albian 
II— III, locality 171, Mlambongwenya Spruit, northern Zululand. 

Description 

The fragment is wholly septate, consisting of three-quarters of a whorl 
only of a juvenile specimen. Coiling is very open, the whorl section depressed, 
reniform, with a very small dorsal impressed zone. Ornament is subdued, and 
consists of single, low prorsiradiate folds on the shell surface. The suture line 
is partially exposed, showing a long E/L saddle with a narrow stem, a large L 
and low L/U saddle. I is trifid (Fig. 53). 

Discussion 

Depressed whorl section, together with the presence of a narrow dorsal 
impressed area place this specimen closest to Pictet & Campiche's species, 
the original illustrations of which are reproduced here as Figure 52D-F; the 
whorl section is more depressed in that species, whilst internal moulds are 
smooth, in consequence of which the authors refer to their specimen as P. aff. 
depressa. In P. astieriana (Fig. 49) the whorl section is more rounded; in 
P. crassecostata Collignon (1963: 8, pi. 243 (fig. 1046)) the whorl section is 
oval with broad, well-differentiated convex ribs; in P. ovalis Collignon (1963: 
8, pi. 243 (fig. 1045)) the whorl section is also oval, with dense, relatively 
prominent flexuous ribs. P. arcuata Collignon (19626: 13, pi. 221 (fig. 958)) is 
an Aptian species based on gently curved fragments with a circular whorl 
section, smooth surface and folds corresponding to the site of individual 
sutures. 



330 ANNALS OF THE SOUTH AFRICAN MUSEUM 

Occurrence 

Pictetia depressa is best known from the Lower Albian of western Europe 
and Transcaspia, although also recorded from the Upper Aptian of Madagascar. 

ACKNOWLEDGEMENTS 

We thank Dr H. W. Ball, Dr M. K. Howarth and Mr D. Phillips of the 
British Museum (Natural History), Dr R. Casey and Mr C. J. Wood (Institute 
of Geological Sciences), Dr J. Sornay (Museum d'Histoire Naturelle, Paris), 
Professor T. Matsumoto (Kyushu, Japan), Dr I. Hayami (Tokyo), Professor 
R. A. Reyment (Uppsala) and Professor D. Visser (Pretoria) for allowing us to 
study material in their care and for assistance in many other ways. 

We also thank General M. Collignon (Moirans), Professor Dr J. Wiedmann 
(Tubingen), Mr C. W. Wright and Dr M. R. Cooper (Oxford) for their advice 
and useful discussions. Dr J. A. Jeletzky of the Canadian Geological Survey 
kindly supplied casts of Ammonoceratites crenocostatus and Professor 
R. A. Reyment (Uppsala) provided photographs of the holotype of 
Argonauticeras belliseptatum for this publication. The help of the staff of the 
Geological Collections, Oxford University Museum ; Geological Survey of South 
Africa, Pretoria; and South African Museum, Cape Town, is gratefully 
acknowledged, as is the financial support of the Trustees of the Sir Henry 
Strakosch Bequest, the Royal Society, and the Natural Environment Research 
Council. 

Thanks are due to the Director, Geological Survey of South Africa, 
Pretoria, for the loan of part of the material described. 



REFERENCES 

Anderson, F. M. 1902. Cretaceous deposits of the Pacific Coast. Proa. Calif. Acad. Sci. 

2: 1-132. 
Anderson, F. M. 1938. Lower Cretaceous deposits in California and Oregon. Spec. Pap. 

geol. Soc. Am. 16: 1-339. 
Anthula, D. J. 1899. Ueber die Kreidefossilien des Kaukasus. Beitr. Palaont. Geol. Ost.-Ung. 

12: 53-159. 
Arkell, W. J. & Wright, C. W. 1957. In: Moore, R. C, ed. Treatise on Invertebrate 

Paleontology Part L, Mollusca 4. Cephalopoda, Ammonoidea. Kansas and New York: 

Geological Society of America & University of Kansas Press. 
Beznosov, N. V. 1958. Jurassic ammonites of the northern Caucasus and Crimea. Phylloceratina 

and Lytoceratina. Leningrad. (In Russian.) 
Bowdich, T. E. 1822. Elements of conchology including the fossil genera of the animals, I 

Univalves. London and Paris. 
Breistroffer, M. 1936. In: Besairie, H. Recherches geologiques a Madagascar. Premiere 

suite. La geologie du Nord-Ouest. Mem. Acad, malgache 21: 9-259. 
Breistroffer, M. & De Villoutreys, O. 1953. Les Ammonites albiennes de Peille (Alpes- 

Maritimes). Trav. Lab. Geol. Univ. Grenoble 30 : 69-74. 
Buckman, S. S. 1905. On certain genera and species of Lytoceratidae. Q. Jl geol. Soc. Lond. 

61: 142-154. 
Buckman, S. S. 1909-1930. Yorkshire type ammonites, London. 



CRETACEOUS FAUNAS FROM SOUTH AFRICA 331 

Casey, R. C. 1960. The ammonoidea of the Lower Greensand. Palaeontogr. Soc. {Monogr.) 

113: 1-44. 
Chenu, J. G. 1859. Manuel de Conchyliologie et de Paleontologie conchyliologique 1. Paris. 
Collignon, M. 1949<3. Faune neocomienne des couches a Crioceras de Belohasifaka (cercle 

de Sitampiky). Annls geol Serv. Mines Madagascar 15: 53-83. 
Collignon, M. 1949&. La faune albienne d'Ambarimaninga. Annls geol. Serv. Mines Mada- 
gascar 16: 1-128. 
Collignon, M. 1950. L'Albien de Mokaraha. Annls geol. Serv. Mines Madagascar 17: 57-85. 
Collignon, M. 1956. Ammonites neocretacees du Menabe (Madagascar). IV.— Les Phyllo- 

ceratidae. V.— Les Gaudryceratidae. VI. — Les Tetragonitidae. Annls geol. Serv. Mines 

Madagascar 23: 1-106. 
Collignon, M. 1962a. Atlas des fossiles caracteristiques de Madagascar {Ammonites) VIII 

(Berriasien, Valanginien, Hauterivien, Barremien). Tananarive: Service Geologique. 
Collignon, M. 19626. Atlas des fossiles caracteristiques de Madagascar {Ammonites) IX 

{Aptien). Tananarive : Service Geologique. 
Collingnon, M. 1963. Atlas des fossiles caracteristiques de Madagascar {Ammonites) X 

{Albien). Tananarive: Service Geologique. 
Crick, G. C. 1907. The Cephalopoda from the tributaries of the Manuan Creek, Zululand. 

Rep. geol. Surv. Natal Zululand 3: 235-249. 
Dimitrova, N. 1967. The fossils of Bulgaria. IV. Lower Cretaceous, Cephalopoda {Nautiloidea 

and Ammonoidea). Bulgarian Academy of Sciences. (In Bulgarian.) 
Douville, H. 1916. Les terrains secondaires dans le Massif du Moghara a l'Est de l'isthme 

de Suez, Paleontologie, Mem. Acad. Sci. 1916: 1-184. 
Drushchitza, V. V. 1953. Observations on the systematics of Lower Cretaceous Ammonites. 

Bjull. Moskovsk. obs. ispit. prirody, N.S. 58: 88-89. (In Russian.) 
Drushchitza, V. V. 1956. Lower Cretaceous ammonites from the Crimea and northern Caucasus. 

Moscow: Moscow University. (In Russian.) 
Drushchitza, V. V. & Kudryavtseva, M. P. 1960. Atlas of Lower Cretaceous faunas of the 

northern Caucasus and Crimea. Moscow: Moscow University. (In Russian.) 
Eristavi, M. S. 1955. Fauna of the Lower Cretaceous of Georgia. Monogr. Inst. geol. miner., 

Akad. Nauk Gruzin S.S.R. 6: 116-117. (In Russian.) 
Eristavi, M. S. 1957. A comparison of the Lower Cretaceous deposits of Georgia and the 

Crimea. Monogr. Inst. geol. miner., Akad. Nauk. Gruzin S.S.R. unnumbered monogr.: 

1-82 (In Russian.) 
Fallot, P. 1920. Observations sur diverses especes du Gargasien bathyal alpin et en particulier 

sur la faune de Blieux. In: Kilian, W. et al. Contribution a l'etude des faunes Paleo- 

cretacees du Sud-est de la France. Mem. Serv. Carte geol. det. Fr. : 229-266. 
Forster, R. 1975. Die geologische Entwicklung von Sud-Mozambique seit der Unterkreide 

und die Ammoniten-Fauna von Unterkreide und Cenoman. Geol. Jb. (B) 12: 1-324. 
Haug, E. 1889. Beitrag zur Kenntniss der oberneocomen Ammoniten Fauna der Puezalpe 

bei Corvara (Sudtirol). Beitr. Paldont. Geol. Ost.-Ung. 7: 193-231. 
Haughton, S. H. 1936.. Account of the geology of the Cretaceous beds, and a preliminary 

analysis of the associated ammonite fauna. In: Rennie, J. V. L. Lower Cretaceous 

lamellibranchia from northern Zululand. Ann. S. Afr. Mus. 21 : 283-297. 
Hyatt, A. 1900. Cephalopoda. In: Zittel, K. A. von. 1896-1900. Textbook of Palaeontology, 

translated by C. R. Eastman: 502-604. London: Macmillan. 
Jacob, C. 1907. Etudes paleontologiques et stratigraphiques sur la partie moyenne des terrains 

cretacees dans les alpes francaises. Trav. Lab. Geol. Univ. Grenoble 8: 280-590. 
Jacob, C. 1908. Etudes sur quelques ammonites du Cretace moyen. Mem. Soc. geol. Fr. 

Paleont. 38: 1-64. 
Jeannet, A. 1908. Une ammonite nouvelle de l'Albien du Jura, Lytoceras sp. aff. mahadeva 

Stoliczka. Bull. Labs. Geol. Geogr. phys. Miner. Univ. Lausanne 9: 1-19. 
Karakasch, N. I. 1907. The Lower Cretaceous of the Crimea and its fauna. Trudy imp. 

S-peterb. Obshch. Estest. 32: 1-482. (In Russian.) 
Kennedy, W. J. & Klinger, H. C. 1975. Cretaceous faunas from Zululand and Natal, South 

Africa. Introduction, Stratigraphy. Bull. Br. Mus. nat. Hist. {Geol.) 25: 265-315. 
Kilian, W. 1892. Sur quelques Ammonites appartenant au Museum d'Histoire naturelle de 

Lyon. Archs Mus. Hist. nat. Lyon 5:8. 



332 ANNALS OF THE SOUTH AFRICAN MUSEUM 

Kilian, W. & Reboul, P. 1915. I. La faune de l'Aptien inferieur des environs de Montelimar 

(Drome). II. Sur quelques ammonites de l'Hauterivien de la Begude (Basses-Alpes). 

In: Kilian, W. et al. Contribution a l'etude des Cephalopodes paleocretaces du sud-est 

de la France. Mem. Serv. Carte geol. det. Fr.: 1-228. 
Kossmat, F. 1895-1898. Untersuchungen iiber die Siidindische Kreideformation. Beitr. 

Palaont. Geol Ost.-Ung. 9 (1895): 97-203; 10 (1897): 1-46; 11 (1898): 89-152. 
Kotetichvili, L. Y. 1970. Stratigrafia i fauna Colchiditovogo i smechnich horizontov zapadnoy 

gruzii. Trudy geol. Inst. Tbilisi n.s. 24: 1-115. (In Russian.) 
Krenkel, E. 1910. Die untere Kreide von Deutsch-Ostafrika. Beitr. Palaont. Geol. Ost.-Ung. 

23: 201-250. 
Ksiazkiewicz, M. 1938. La fauna des Neocomien superieur de Lanckorona. C.R. Comm. 

Phys. Acad. Polonaise des Lettres 62: 223-261. 
Kullmann, J. & Wiedmann, J. 1970. Significance of sutures in phylogeny of Ammonoidea. 

Paleont. Contr. Univ. Kans. 47: 1-32. 
Kulzhinskaya-Voronets, N. S. 1933. Predstaviteli semejstava Lytoceratidae Neumayr 

emend. Zittel iz niznemelovych otlozenij Kryma. Trudy Vsesoyuznogo geologo- 

razbedoynogo obeduneuya N.K.T. S.S.S.R. 241: 1-38. 
Lamarck, J. B. P. A. de Monet, Chevalier de. 1815-1822. Histoire naturelle des animaux sans 

vertebres. 7 vols. Paris. 
McLearn, F. H. 1972. Ammonoids of the Lower Cretaceous Sandstone Member of the Haida 

Formation, Skidegate Inlet, Queen Charlotte Islands, Western British Columbia. Bull. 

geol. Surv. Can. 188: 1-78. 
Matheron, P. 1878-1880. Recherches paleontologiques dans le Midi de la France. 7 livraisons, 

pis (incomplete). Marseilles. 
Matsumoto, T. 1954. The Cretaceous System in the Japanese Islands. Tokyo: Japanese Society 

for the Promotion of Science. 
Murphy, M. A. 1975. Paleontology and stratigraphy of the Lower Chickabally Mudstone 

(Barremian-Aptian) in the Ono Quadrangle, Northern California. Univ. Calif. Pubis 

geol. Sci. 113: 1-52. 
Nagy, I. Z. 1967. Unterkretazische Cephalopoden aus dem Gerecse-Gebirge 1. Annls hist.-nat. 

Mus. natn. hung. 59: 53-79. 
Neumayr, M. 1875. Die Ammoniten der Kreide und die Systematik der Ammonitiden. 

Z. dt. geol. Ges. 27: 854-942. 
Oppel, A. 1865. Die tithonische Etage. Z. dt. geol. Ges. 17: 535-558. 
Orbigny, A. d'. 1840-1842. Paleontologie francaise. Terrains cretaces. 1. Cephalopodes. 

Paris: Masson. 
Orbigny, A. d'. 1850. Prodrome de Paleontologie stratigraphique universelle des animaux, 

mollusques et rayonnes, 11. Paris: Masson. 
Orlov, Y. U., ed. 1958. Osnovy Paleontologii VI. Mollusca-Cephalopoda II. Moscow: 

Gosudarstvennoe Nauchno-Tekhnicheskoe Izdatel'stvo Literatury po Geologii i Okhrane 

Nedr. (In Russian.) 
Pervinquiere, L. 1907. Etudes de paleontologie tunisienne. 1. Cephalopodes des terrains 

secondaires. Mem. Carte geol. Tunis.: 1-483. 
Petkovic, V. K. 1921. O barremskom katu na Grebenu. Glas Srpskje Kraljveske Akad. 95: 

35-78. (In Serbian). 
Pictet, F. J. & Campiche, G. 1858-1864. Materiaux pour la Paleontologie Suisse. Description 

des fossiles du terrain Cretace des environs de Ste. Croix II, no. 2; III, no. 2. Geneva. 
Rafinesque-Schmaltz, C. S. 1815. Analyse de la Nature, ou tableaux d'Univers des corps 

organisees. Palermo. 
Rouchadze, J. 1933. Les ammonites aptiennes de la Georgie occidentale. Bull. Inst. geol. 

Georgie 3: 165-273. 
Sarasin, C. & Schondelmayer, C. 1901-1902. Etude monographique des ammonites du 

cretacique inferieur de Chatel-Saint-Denis. Abh. schweiz. palaont. Ges. 28 (1901): 1-91; 

29(1902): 95-195. 
Schindewolf, O. 1960. Studien zur Stammesgeschichte der Ammoniten. Part 1. Abh. math.- 

naturw. Kl. Akad. Wiss. Mainz 1960 (10): 635-743 (1-109). 
Simionescu, J. 1898. Studii geologice si paleontologice din Carpatii sudici i Studii geologice 



CRETACEOUS FAUNAS FROM SOUTH AFRICA 333 

asupra basenului Dimboviciorei. ii— Fauna neoconvana din basenul Dimboviciorei. 

Publnile Fond. Vasilie Adamachi 1 : 61-167. 
Simionescu, J. 1900. Notes sur quelques Ammonites du Neocomien Francaise. Trav. Lab. 

Geol. Univ. Grenoble 5'. 1-17. 
Somogyi, K. 1914. Das Neocom des Gerecsegebirges. Annls hist.-nat. Mus. natn. hung. 22: 

277-345. 
Sowerby, J. 1818-1821. The Mineral conchology of Great Britain; or coloured figures and 

descriptions of those remains of testaceous animals or shells, which 'have been preserved 

at various times and depths in the earth. Vol . 3 , pis 204-306 (pis 204-22 1 : 1818; pis 223-253 : 

1819; pis 254-271: 1820; pis 272-306: 1821). London. 
Spath, L. F. 1923-1943. A Monograph of the Ammonoidea of the Gault. Palaeontogr. Soc. 

(Monogr.): 1-787. 
Spath, L. F. 1927. Revision of the Jurassic Cephalopod fauna of Kachh (Cutch). Mem. geol. 

Surv. India Palaeont. indica n.s. 11: 1-71. 
Spath, L. F. 1939. The Cephalopoda of the Neocomian Belemnite Beds of the Salt Range. 

Mem. geol. Surv. India Palaeont. indica n.s. 25: 1-154. 
Stoliczka, F. 1863-1866. The fossil Cephalopoda of the Cretaceous rocks of southern India. 

Mem. geol. Surv. India Palaeont : indica 1: 41-56(1863); 2-5: 47-106 (1864); 6-9: 107-154 

(1865); 10-13: 155-216 (1866). 
Suess, E. 1865. Ober Ammoniten. Sitz.-Ber. k.k. Akad. Wiss. 52: 71-89; 305-322. 
Thomel, G. 1968. A propos de Y Ammonoceratites (Argonauticeras) depereti (Kilian) 

du Gargasien des Basses-Alpes orientales. Bull. Soc. geol. Fr. (7) 10: 684-687. 
Uhlig, V. 1883. Die Cephalopoden der Wernsdorfer Schichten. Denkschr. Akad. Wiss., Wien 

46(2): 127-290. 
Vasicek, Z. 1972. Ammonoidea of the Tesin-Hradiste Formation (Lower Cretaceous) in the 

Moravskoslzske Beskydy Mountains. Ustfedniho ustavu geologickeho 38: 1-103. 
Wedekind, R. 1916. Ober Lobus, Suturallobus und Inzision. Zentbl. Miner. Geol. Paldont. 

1916: 185-195. 
Whiteaves, J. F. 1876. On some invertebrates from the coal-bearing rocks of the Queen 

Charlotte Islands. Mesozoic Fossils 1 (1): 1-92. Ottawa: Geological Survey of Canada. 
Whiteaves, J. F. 1884. On the fossils of the coal-bearing deposits of the Queen Charlotte 

Islands collected by Dr G. M. Dawson in 1878. Mesozoic Fossils 1 (3): 191-262. Ottawa: 

Geological Survey of Canada. 
Whiteaves, J. F. 1900. On some additional or imperfectly understood fossils from the 

Cretaceous Rocks of the Queen Charlotte Islands, with a revised list of the species from 

these rocks. Mesozoic Fossils 1 (4): 263-308. 
Wiedmann, J. 1962. Unterkreide-ammoniten von Mallorca I. Liefr. Lytoceratina, Aptychi. 

Abh. math.-naturw. Kl. Akad. Wiss. Mainz 1962 (1): 1-148. 
Wiedmann, J. & Dieni, I. 1968. Die Kreide Sardiniens und ihre Cephalopoden. Palaeontogr. 

ital. 64: 1-171. 
Yabe, H. 1903. Cretaceous Cephalopoda from Hokkaido. Part 1. /. Coll. Sci. imp. Univ. 

Tokyo 18: 1-55. 
Zittel, K. A. von. 1868. Die Cephalopoden der Stramberger Schichten. Palaeont. Mitt. Mus. k. 

bayer. St. 2: 1-118. 
Zittel, K. A. von. 1884. Handbuch der Palaeontologie. (Palaeozoologie). i Abt., ii Band, Lief. 

iii. Cephalopoda. Munich & Leipzig: Oldenbourg. 
Zwierzycki, J. 1914. Die Cephalopoden-fauna der Tendaguru-schichten in Deutsch-Ostafrika. 

Arch. Biontol. 3 : 7-96. 



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Names of new taxa, combinations, synonyms, etc., when used for the first time, must be 
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Family Nuculanidae 

Nuculana (Lembulus) bicuspidata (Gould, 1 845) 

Figs 14-1 5 A 
Nucula {Leda) bicuspidata Gould, 1845: 37. 
Leda plicifera A. Adams, 1856: 50. 

Laeda bicuspidata Hanley, 1859: 118, pi. 228 (fig. 73). Sowerby, 1871 : pi. 2 (fig. 8a-b). 
Nucula largiilicrti Philippi, 1861 : 87. 
Leda bicuspidata: Nickles, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9. 

Note punctuation in the above example: 

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SAM-A13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King's Beach 
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Name of new genus or species is not to be included in the title : it should be included in the 

abstract, counter *o Recommendation 23 of the Code, to meet the requirements of 

Biological Abstracts. 



WILLIAM JAMES KENNEDY 

& 
HERBERT CHRISTIAN KLINGER 

CRETACEOUS FAUNAS FROM ZULULAND 

AND NATAL, SOUTH AFRICA 
THE AMMONITE FAMILY LYTOCERATIDAE 

NEUMAYR, 1875 



VOLUME 74 PART 13 MARCH 1978 



ISSN 0303-2515 



MUS. COMP, ZOOL. 
LIBRARY 

J UN 2 1978 

HARVARD 
UNIVERSITY 



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Examples (note capitalization and punctuation) 

Bullough, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan. 

Fischer, P.-H. 1948. Donnees sur la resistance et de le vitalite des mollusques. /. Conch., Paris 88: 100-140. 

Fischer, P.-H., Duval, M. & Raffy, A. 1933. Etudes sur les echanges respiratoires des littorines. Archs 

Zool. exp. gen. 74: 627-634. 
Kohn, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon. 

Ann. Mag. nat. Hist. (13) 2: 309-320. 
Kohn, A. J. 19606. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean. 

Bull. Bingham oceanogr. Coll. 17 (4): 1-51. 
Thtele, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. In: Schultze, L. Zoologische 

und anthropologische Ergebnisse einer Forschungsreise im west lichen und zentralen Sud-Afrika 4 : 269—270. 

Jena: Fischer. Denkschr. med.-naturw. Ges. Jena 16: 269-270. 

(continued inside back cover) 



ANNALS OF THE SOUTH AFRICAN MUSEUM 
ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM 



Volume 74 Band 
March 1978 Maart 
Part 13 Deel 




RE-EXAMINATION OF ARCHAEICHNIUM, 
A FOSSIL FROM THE NAMA GROUP 



By 



MARTIN F. GLAESSNER 



Cape Town Kaapstad 



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RE-EXAMINATION OF ARCHAEICHNIUM, A FOSSIL FROM 

THE NAMA GROUP 

By 

Martin F. Glaessner 
University of Adelaide, South Australia 

(With 3 figures) 
[MS. accepted 6 September 1977] 

ABSTRACT 

The fossils described by Haughton in 1960 as Archaeocyatha from the Nama Group were 
later, after the study of plaster casts, considered as worm tubes or worm burrows by the 
present author and named Archaeichnium haughtoni. The type specimens have now been 
re-examined. They consist of agglutinated sand grains and are similar to tubes made actively 
by living polychaete worms. They are not closely similar to the partly calcareous tubes of 
Cloudina or to the distinctively structured Volborthella. In the absence of any taxonomically 
significant characters of tube construction the systematic position of the originator of these 
fossils remains uncertain. 

CONTENTS 

PAGE 

Introduction 335 

New observations 338 

Interpretation 338 

Conclusions 341 

Acknowledgements 342 

References . . 342 

INTRODUCTION 

Haughton (1960) described what he considered 'An archaeocyathid from 
the Nama System', from two specimens (housed in the South African Museum) 
of a quartzitic sandstone on Griindoorn near the Ham River, 'a typical member 
of the Kuibis beds'. Germs later re-examined the stratigraphy of the Nama 
Group. From his brief description of the eastern area it appears that Haughton's 
fossils came from the Upper Clastic Member of the Nababis Formation, probably 
corresponding to the Upper Clastic Member of the Kuibis Formation in the west 
(Germs 1972a: 19, figs 14, 22). The interpretation of these fossils hinges on their 
mode of fossilization. It is therefore necessary to summarize Haughton's views 
on this matter. He considered them as moulds on bedding planes and noted 
correctly that some of the supposed moulds are 'marked off' by a bordering 
depression and that some are lying parallel to each other, as if deposited by 
current action. They are described as elongated tubular cones with circular cross- 
sections. 'The more complete individuals have a pointed lower end.' In two 

335 
Ann. S. Afr. Mus. 74 (13), 1977: 335-342, 3 figs. 



336 ANNALS OF THE SOUTH AFRICAN MUSEUM 

instances, 'two individuals appear to have been joined at this pointed end'. 
Haughton noted also the presence of a number of elongate (longitudinal) 
'pustulate ridges' and of an individual 'mould' appearing as an approximately 
circular ridge which 'can be interpreted as an infilling of the intervallum or space 
between the internal and external walls of a cup, or as a replacement of a single 
thick exoskeleton'. The second alternative was abandoned without further com- 
ment and the first one adopted. Thus the longitudinal section (as seen below the 
centre of Fig. 1) was interpreted as showing an infilling of a central cavity sur- 
rounded by concentric inner and outer walls which are now indicated by 
depressions. The solid specimens being considered as internal moulds, the present 
longitudinal ridges on their outer surfaces were thought to separate what would 
have been broad, rounded ridges on the inner surfaces of the vanished outer 
walls. If pustules are now present, they would have originated as rows 'of evenly 
spaced pores' in longitudinal channels. This reconstruction of the fossils from 
their supposed 'moulds' led Haughton without much difficulty to his interpreta- 
tion as Archaeocyatha. 

The obvious significance of Haughton's classification of his fossils for the 
dating of the Nama Group led to their re-examination (Glaessner 1963). Those 
studies were based on plaster casts of the originals. It appeared that the fossils 
represented infillings of cemented tubes made, probably, by worm-like animals 
which incompletely agglutinated the sediment surrounding their burrows. The 
tubes were considered as generally cylindrical but they appeared to be conical 
where they entered the rock matrix. The specimen (below the centre in Fig. 1, 
marked M in Glaessner 1963, pi. 3 (fig. 1)) which Haughton saw as two indivi- 
duals joined at their pointed ends, represents one tube dipping from the bedding 
plane into the rock matrix and re-emerging with a 90° change in direction. The 
other specimen, marked with an arrow on Haughton's Plate 4 (L in Glaessner 
1963, pi. 3 (fig. 1)), shows a similar change of direction. The number of longitu- 
dinal ridges was correctly described by Haughton as ten to twelve on each 
individual. Five to six ridges can be seen on one side but the supposed pustules 
are sand grains. The ridges are also visible on several fragmentary specimens but 
appear only faintly at the ends of the largest specimen. The longitudinal ridges 
on the outer surfaces of the agglutinated tubes and the transverse layering of the 
1 mm thick walls appear to be diagnostic for these fossils. None of the distinctive 
characters of the Archaeocyatha can be seen in them. There are no pointed cones, 
no traces of a former calcareous double wall, no septa or other intervallum 
structures. There is no evidence of an intervallum. 

The description of the partly agglutinated tubes of Cloudina by Germs 
(1972a, 19726) and recent studies of this genus and of the fully agglutinated 
Volborthella (Glaessner 1976) suggested that a re-examination of Archaeichnium 
would be desirable, not because they could still be considered as Archaeocyatha 
or as identical with Cloudina, as subsequently suggested by Kaever & Richter 
(1976: 30), but because more is now known about early tube-building worms. 
The observations of the morphology of these fossils made previously on plaster 



RE-EXAMINATION OF ARCHAEICHNIUM, A FOSSIL FROM THE NAMA GROUP 337 




Fig. 1. Archaeichnium haughtoni Glaessner, 1963, on a slab of quartzite from the Nama Group 
(Late Precambrian). Slab B, SAM-K4812. The holotype is the specimen below the centre of 

the figure. Natural size. 



338 ANNALS OF THE SOUTH AFRICAN MUSEUM 

casts have been supplemented by examination of the original specimens. One of 
them is marked Slab A, SAM-K4813, referred to correctly as Slab A by Haugh- 
ton (1960) on pages 57 and 58, but as Slab B on plate 3 (true magnification x 0,9, 
not 'above natural size'). The other is marked Slab B, SAM-K4812, which is 
correctly referred to by Haughton on pages 57 and 58 and on plate 5, but as 
Slab A on plate 4. The true magnification of this plate is about x 1,5, not x 5/8. 
Part of a plaster cast of Slab A was previously figured (Glaessner 1 963) as plate 3, 
figure 2, and part of Slab B as plate 3, figure 1 and they are now shown as 
Figures 1 (Slab B) and 2 (Slab A). 

NEW OBSERVATIONS 

The fossils consist of firmly cemented sand grains, mostly quartz. Although 
the slabs could not be cut and without sections no measurements could be made, 
the grains forming the tubes appear to be slightly coarser and more uniform than 
the matrix. They must have been agglutinated prior to fossilization as they are 
mostly sharply delimited from the matrix and do not grade into it on the bedding 
planes. There is no indication of any previous presence of thin 'walls' between 
the matrix and the fossils. The rock slabs consist of a slightly glassy quartzite. 
Diagenetic alteration of a sandy layer has fused the tubes with the sediment 
represented by the slabs, leaving them projecting in sharp semirelief from one 
bedding plane. On the analogy of similarly preserved Australian fossils collected 
in situ, this bedding plane was probably underlain by a silt or clay layer. The 
preservation is likely to be in hyporelief. On Slab A (Fig. 2) and possibly also 
on Slab B (Fig. 1) where numbers are smaller, the tubes show (as Haughton had 
recognized) a preferred orientation, indicating some current sorting. This is 
demonstrated by Figure 3. Sizes of the tubular fossils range from 1 mm to almost 
5 mm in diameter. There is some flattening and some apparent twisting of tubes. 
Some are cylindrical, some appear to be barrel-shaped, some were probably 
conical and a few, including one near the bottom of Figure 1 (L in pi. 3 (fig. 1) 
in Glaessner 1963) thin abruptly at one end to form a long, thin, twisted tail. 
Irregular forms are more common than geometrically regular ones. Some ends 
are sharp, others fade out in the matrix. Few if any clean breaks which would 
indicate brittleness are visible and it is concluded that the tubes were compres- 
sible and flexible, particularly where their walls were thin. 

INTERPRETATION 

In 1963 it was demonstrated that the fossils could not be considered as 
Archaeocyatha. What are these fossils? Specimen M (Fig. 1, below centre) was 
selected as the holotype of Archaeichnium haughtoni because it shows the most 
distinctive characters of the new monotypic genus which are seen less clearly 
among the other fossils on the two slabs. The holotype specimen can be followed 
in appropriate lighting conditions over a length of 70 mm, almost to the edge of 



RE-EXAMINATION OF ARCHAEICHNIUM, A FOSSIL FROM THE NAMA GROUP 339 




Fig. 2. Archaeichnium haughtoni Glaessner, 1963. Slab A, SAM-K4813. (The lobe projecting 
from the lower part of the bedding plane may be a bioturbated load cast.) Natural size. 



340 



ANNALS OF THE SOUTH AFRICAN MUSEUM 





23 


23 












7 










4 






4 




3 




3 




2 


2 


2 



90 80 60 40 20 



0^ 



20 40 60 8090 



Fig. 3. Histogram of positions of 73 individual fossils on Slab A showing deviation of long 

axis from modal direction. 



the slab. Its maximum width is almost 5 mm. Its wall is about 1 mm thick and 
shows pronounced transverse layering of sand grains. The centre of the tube is 
filled with sand grains which show a much fainter and less regular transverse 
fabric. The outer surface of the tube is marked with fine longitudinal ribs, but 
they are visible in this specimen only faintly near the ends of the tube, which is 
either smoothed by weathering or split open for most of its length. At the end 
nearer the edge of the slab the ribs are preserved as impressions in the matrix, 
hence they are an external sculpture, not part of an internal mould. In the other 
large specimen (bottom of Fig. 1, L in pi. 3 (fig. 1) in Glaessner 1963) the 
external ribs are well preserved and straight but its internal structure is not 
revealed. This tube is 35 mm long and up to 4,5 mm wide. At one end it is 
narrowed and appears twisted. Several other specimens show some of these 
characters such as external longitudinal ribbing or transverse wall fabric. Their 
size ranges fit those of the larger specimens and the indications of original 
flexibility are similar. On these grounds the author sees no reason to distinguisl 
any specimens taxonomically from the typical ones. 

It is difficult to decide whether these fossils are burrows in the strict sens* 
of the word. There is no doubt that they were built by a worm-shaped animal 
from sand grains selected and agglutinated by some organic cement which 
allowed them some flexibility. The clear separation between the tube walls and 
the matrix brought out by weathering of the bedding plane indicates that the 
composition of the walls was different from that of the matrix. The wall material 
appears to have been actively selected by the animal. There is no evidence for 
calcareous walls having been dissolved during fossilization. The selection of 
grains and their arrangement so as to form external and possibly strengthening 
ridges shows active building from transported grains, rather than passive cemen- 



RE-EXAMINATION OF ARCHAEICHNIUM, A FOSSIL FROM THE NAMA GROUP 341 

tation of a burrow wall from grains of matrix in situ. Faint transverse texture 
of the infilling of the tube in the holotype specimen may indicate active filling, 
but there is no confirmation from other specimens. It is likely that the tubes were 
made by organisms burrowing in or living on the sediment and constructing 
agglutinated walls from sand grains. It is also probable that subsequent current 
action as indicated particularly by the orientation of burrow fragments on 
Slab A washed the weakly cemented burrows or tubes out of a layer of sediment 
and distributed them over a bedding plane. If this were the case, it raises the 
semantic point whether we are dealing with trace fossils in the strict sense of the 
word or whether worm tubes agglutinated from sand grains with organic cement 
are properly considered as body fossils, but this is not a matter of substantial 
importance. The Slabs A and B are so similar that they may well represent not 
very distant parts of the same bedding plane, yet one (A) indicates current action 
while the other (B) may be thought to have at least the two curved, long tubes 
preserved in situ. These questions cannot be solved without further field studies 
and discovery of material in situ. 

CONCLUSIONS 

Archaeichnium haughtoni is a name given to tubular fossils with walls 
agglutinated from small detrital quartz grains. The best preserved of these tubes 
show external, longitudinal, straight, fine ribs and a transverse layering of the 
grains forming the wall. They may have been actively infilled. Most of the tubes 
seem to be fragments washed out of the original sediment within or from which 
they were built. They appear to have been redeposited on bedding planes where 
they were subsequently buried by layers of sand and diagenetically fused with it. 
There is no significant resemblance between this fossil and others from the Nama 
Group or indeed from anywhere else. After this study had commenced, Kaever 
& Richter (1976) described Buschmannia roeringi from the Buschmannsklippe 
Formation of the Nama Group as Archaeocyatha. The author was able to 
re-examine this fossil. Its resemblance to Praesolenopora Tsao & Zhao, 1972, 
from the Late Precambrian (Upper Sinian) of China (see Tsao & Liang 1974) 
was noted and it was concluded that it is probably a calcareous alga. 
Archaeichnium differs from Cloudina Germs from the Nama Group (Germs 
1972a, 19726). This genus builds partly calcareous, secreted tubes with only 
minor, external agglutination of sand grains and transverse sculpture. 'The tube 
walls seem to have contained organic material. Their distortions suggest some 
flexibility . . . reminiscent of some agglutinating foraminifera with walls of 
quartz or carbonate grains and much organic cement' (Glaessner 1976: 266). 
Cloudina is generally placed in the Order Cribricyathida, most of which are of 
early Cambrian age. Their morphological characters make probable their 
'possible relation to the serpulids' (Germs 19726: 753) and suggest that they are 
'best classified as an extinct Order of the Class Polychaeta' (Glaessner 1976: 268). 
Archaeichnium differs from Volborthella which is built from invaginated cones 



342 ANNALS OF THE SOUTH AFRICAN MUSEUM 

of agglutinated sandy material round a central tube in distinctly sloping layers, 
and from the similar but predominantly calcareous Salterella. This mode of tube 
construction is similar in some detail to that used by living Sabellariidae. 
Volborthella can be considered as the product of polychaete annelid worms. 
While the tubes and the inferred tube-building activities of Archaeichnium are 
comparable and compatible with those of polychaete worms of the Order Tere- 
bellida (and probably others), there is as yet nothing sufficiently specific in the 
known morphological characters of this fossil to support its definite systematic 
assignment to the Annelida Polychaeta. It can only be tentatively attached to 
them and must remain for the present incertae sedis. No occurrences of this fossil 
other than that at its type locality, in strata considered to be of Late Precambrian 
age, are known. 

ACKNOWLEDGEMENTS 

I wish to thank the authorities of the South African Museum for the loan 
of the type material for this study; the support of this loan by Mr P. J. Rossouw, 
South African Geological Survey, Pretoria, is gratefully acknowledged. Through 
the kind co-operation of Prof. M. Kaever I was able to re-examine Buschmannia 
roeringi. Mr Richard Barrett, Department of Geology, University of Adelaide, 
took the photographs of the rock specimens. 

REFERENCES 

Germs, G. J. B. 1972a. The stratigraphy and paleontology of the lower Nama Group, South 

West Africa. Bull. Chamber Mines Precambr. Res. Unit 12. 
Germs, G. J. B. \912b. New shelly fossils from the Nama Group, South West Africa. Am. J. 

Sci. Ill: 752-761. 
Glaessner, M. F. 1963. Zur Kenntnis der Nama-Fossilien Sudwest-Afrikas. Annln naturh. 

Mus. Wien 66: 133-120. 
Glaessner, M. F. 1976. Early Phanerozoic annelid worms and their geological and biological 

significance. /. geol. Soc. Lond. 132: 259-275. 
Haughton, S. H. 1960. An archaeocyathid from the Nama System. Trans. R. Soc. S. Afr. 36: 

57-59. 
Kaever, M. & Richter, P. 1976. Buschmannia roeringi n. gen. n. sp. (Archaeocyatha) aus dei 

Nama-Gruppe Siidwestafrikas. Palaont. Z. 50: 27-33. 
Tsao, Rui-Chi & Liang, Yu-Zhou. 1974. On the classification and correlation of the Sinian 

System in China, based on a study of algae and stromatolites. Mem. Nanking Inst. Geol. 

Pal. 5: 1-26. 



6. SYSTEMATIC papers must conform to the International code of zoological nomenclature 
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Names of new taxa, combinations, synonyms, etc., when used for the first time, must be 
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Synonymy arrangement should be according to chronology of names, i.e. all published 
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Family Nuculanidae 

Nuculana (Lembulus) bicuspidata (Gould, 1 845) 

Figs 14-15A 
Pellicula (Leda) bicuspidata Gould, 1845: 37. 
Leda plicifera A. Adams, 1856: 50. 

Laeda bicuspidata Hanley, 1859: 118, pi. 228 (fig. 73). Sowerby, 1871: pi. 2 (fig. 8a-b). 
Nucula largillierti Philippi, 1861 : 87. 
Leda bicuspidata: Nickles, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9. 

Note punctuation in the above example: 

comma separates author's name and year 

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In describing new species, one specimen must be designated as the holotype; other speci- 
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Holotype ' 

SAM-A 13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King's Beach 
Port Elizabeth (33°51'S 25°39'E), collected by A. Smith, 15 January 1973. 

Note standard form of writing South African Museum registration numbers and date. 

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Capital initial letters 

(a) The Figures, Maps and Tables of the paper when referred to in the text 

e.g. '. . . the Figure depicting C. namacolus . . .'; '. . . in C. namacolus (Fig. 10) . . .' 

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Biological Abstracts. 



MARTIN F. GLAESSNER 

RE-EXAMINATION OF ARCHAEICHNIUM, 
A FOSSIL FROM THE NAMA GROUP 



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